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Journal of 

Hymenoptera 
Research 



Volume 10, Number 1 April 2001 

ISSN #1070-9428 
CONTENTS 

BAYLISS, P. S. and D. J. BROTHERS. Behaviour and host relationships of DoUchoniutilla 

sycorax (Smith) (Hymenoptera: MutilHdae, Sphecidae) 1 

CARVALHO, G. A. The number of sex alleles (CSD) in a bee population and its practical 

importance (Hymenoptera: Apidae) 10 

GESS, S. K. Distribution and ethology of Priscomasaris Gess (Hymenoptera: Vespidae: 

Masarinae: Priscomasarina) in Namibia 16 

GIBSON, G. A. P. The Australian species of Pachyneuron Walker (Hymenoptera: Chalcid- 

oidea: Pteromalidae) 29 

JANJIC, J. and L. PACKER. New descriptions of Halictus (Seladonia) from the New World 

(Hymenoptera: Halictidae) 55 

KIMSEY, L. The new Western Australian tiphiid genus Dythynmis (Hymenoptera: Tiphii- 

dae: Thynninae) 76 

MASON, P G., M. A. ERLANDSON, and B. J. YOUNGS. Effects of parasitism by 
Banchus flavescens (Hymenoptera: Ichneumonidae) and Microplitis mediator (Hy- 
menoptera: Braconidae) on the bertha armyworm, Mamesira configurata 
(Lepidoptera: Noctuidae) 81 

MASSA, B., M. C. RIZZO, and V. CALECA. Nahiral alternative hosts of Eulophidae 
(Hymenoptera: Chalcidoidea) parasitoids of the citrus leafminer Phyllocnistis cit- 
rella Stainton (Lepidoptera: Gracillariidae) in the Mediterranean basin 91 

SHAW, M. R. Interactions between adults of some species of Netelia Gray (Hymenoptera: 

Ichneumonidae: Tryphoninae) and their caterpillar hosts (Lepidoptera) 101 




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This issue was mailed 26 April 2001 



J. HYM. RES. 
Vol. 10(1), 2001, pp. 1-9 

Behaviour and Host Relationships of Dolichomutilla sycorax (Smith) 
(Hymenoptera: Mutillidae, Sphecidae) 

Paul S. Bayliss and Denis J. Brothers 

School of Botany & Zoology, University of Natal (Pietermaritzburg), Private Bag XOl, 
Scottsville, 3209 South Africa; (PSB: Current address: Transvaal Museum, P.O. Box 413, 

Pretoria, 0001 South Africa) 



Abstract. — Detailed biological information for species of Mutillidae is generally lacking. The 
following aspects of the biology of Dolichomutilla sycorax (Smith), based on laboratory observations 
of 10 specimens (99, 16) reared from a single nest of Sceliyhron spirifex (Limiaeus) (Sphecidae), 
are described in detail and discussed: emergence from host nest, activity patterns, mating and 
grooming. The recorded host relations for D. si/cornx are also discussed. 



Successful mating by mutillid wasps 
generally requires only a short time 
(Brothers 1972), and very few observations 
have been recorded. These are important 
in providing information on sex associa- 
tions. Grooming in mutillids has recently 
been described for the first time (Bayliss 
and Brothers 1996), and only in the last 
few years has it been used in systematic 
studies of the Hymenoptera (Basibuyuk 
and Quicke 1999). This paper provides the 
first descriptions of mating and grooming 
behaviour in Dolichomutilla si/corax (Smith) 
and surveys the data on its host relation- 
ships, as well as providing some other in- 
cidental information on the behaviour of 
this species. 

Dolichomutilla si/corax is broadly distrib- 
uted throughout eastern Africa, from Ken- 
ya to South Africa where it is the most 
common species of the genus. Its taxo- 
nomic status as a valid species distinct 
from Dolichomutilla guitieeusis (Fabricius) 
has recently been clarified by Nonveiller 
(1996). Specimens are approximately 9-22 
mm long, with the head and metasoma 
black and the niesosoma deep maroon- 
red. The apterous females have a pair of 
w^hite spots on the second metasomal ter- 
gum and an interrupted broad white band 



on the third tergum; the macropterous 
males are almost identical in coloration, 
unlike for most Mutillidae, and have con- 
spicuously banded wings (Figs. 1-2). Al- 
though Gerstaecker (1857, 1862) first de- 
scribed the male (misidentified as that of 
D. guiueeusis), presumably based on the 
similarity of the sexes, Peringuey (1898) 
was the first to associate the sexes directly, 
having reared both simultaneously from 
the mud nests of Pelopaeus [= Sceliphrou] 
spirifex (Linnaeus) (Hymenoptera, Spheci- 
dae). 

MATERIALS AND METHODS 

Laboratory observations were made 
during April to December 1996 at the Uni- 
versity of Natal, Pietermaritzburg. Live 
adults of D. sycorax were reared from a 
mud nest of S. spirifex collected at the 
Greater St Lucia Wetland Park, Ozabeni 
Section, Lower Mkuze, KwaZulu-Natal, 
South Africa (27°39'S, 32°26'E) on 6-9 
April 1996 by R.M. Miller and J. Kotze. 
They emerged over a period of about 10 
days (starting on 19 May 1996) and were 
kept isolated in petri dishes (diameter 90 
mm, height 20 mm) after emergence. The 
bases of the dishes were lined with paper 
towelling to provide a rough substrate. 



Journal of Hymenoptera Research 




'-i^*^^ 




Figs. 1-2. Dolichomutilla si/cornx. 1, 6, lateral view 
(body length 10 mm). 2, 9, dorsal view (body length 
16 mm). 



Mating was observed by placing two 
adults of opposite sex in the same petri 
dish; their behaviour was recorded using 
a Sony 8mm video camera and low-inten- 
sity cool fibre-optic illumination. The 
specimens were observed for at least 20 
minutes, and if no interaction (including 
stridulation or rubbing of antennae) oc- 
curred between them during that time, 
they were separated for several hours be- 
fore placing them together again. 

A Wild M5 stereo microscope, using 
white light from a desk lamp, was used 
for observations of grooming at irregular 
intervals during the day and at night. 
Such behaviour was noted whenever seen, 
and detailed observations were carried 
out after sprinkling the body with flour. 
Observations (a total of at least 60) in- 
volved nine males and one female of D. 
sycorax that emerged from the mud nest 
and the full repertoire of cleaning activi- 
ties was seen 12 times in seven different 
individuals (6 9, Id). The terminology 



used in describing the grooming behav- 
iour is from Basibuyuk and Quicke (1999). 
After sufficient observations had been 
made, the specimens were released into a 
glass terrarium (288 x 217 X 225 mm, in- 
ternal measurements) with the floor cov- 
ered by fine sand to a depth of 25 mm and 
with several flattish stones to provide hid- 
ing places. Food (a solution of 10% honey 
dissolved in water) and water were pro- 
vided in small glass tubes plugged with 
cotton wool. Most specimens lived for 3- 
10 months, the male surviving for the 
shortest period (8 weeks). 

RESULTS AND DISCUSSION 

Specimens Emerging from Mud Nest of 
Host 

The mud nest comprised 15 more or less 
parallel cells separated by thick walls and 
with the outer walls thickened and rough- 
ened by the addition of extra mud. Within 
16 days after the first recorded emergence, 
one male (12 mm long) and nine females 
(12-15 mm long) of D. sycorax, one speci- 
men of Stilhum cyanurum (Forster) (Chry- 
sididae) and two specimens of Sceliphron 
spirifex (one of each sex) had emerged. In 
addition, there were two cells containing 
host cocoons which produced hundreds of 
specimens of a species of Melittobia (Eu- 
lophidae). The rate of parasitism was thus 
87%. 

Activity Patterns of D. sycorax 

Emergence. — It took approximately 10 
minutes for each individual, using the 
mandibles, to chew its way out of the cell. 
The antennae, followed by the head, first 
emerged through the newly chewed exit 
hole, and the surroundings were scanned. 
Since the forepart of the body is often 
slightly narrower than the posterior part, 
the metasoma was often unable to pass 
through the hole. The process of chewing 
would then be resumed until the hole was 
large enough for the entire body to pass 
through. After emergence, several minutes 



Volume 10, Number 1, 2001 



were spent inspecting the nest, although 
no attempt was made to enter a previous- 
ly vacated cell (n = 3). 

Daily activity. — Because of the artificial 
conditions of the terrarium, it is impossi- 
ble to assume much about daily cycles. A 
female was placed together with the oth- 
ers in the terrarium only after the male 
had mated or interacted with her. The 
only male was kept in the terrarium with 
the niated females and his activity, as with 
the females, was monitored. At night all 
females huddled together under the same 
flat stone, even though there were several 
others of similar shape. The male was sol- 
itary, never resting with the females. The 
male died after 55 days, while the females 
lived for approximately 6 months. One fe- 
male lived for almost 11 months. 

Mating 

Immediately after a male and a female 
were placed together in a petri dish, after 
having been kept in separate vials (n = 5), 
they initially tried to escape by running. 
Whenever the two individuals came into 
contact head-on, both instantly showed 
avoidance or escape reactions by moving 
away in different directions. This is simi- 
lar to Ferguson's (1962) observations on 
Sphaeropthalma (Photopsis) blakeii (Fox) but 
contrary to Brothers' (1972) observations 
on Pseudomethoca frigida (Smith) and Bay- 
liss and Brothers' (1996) observations on 
Tridiolabiodes spp. where neither member 
showed avoidance reactions. As soon as 
the male contacted the female, except 
when head-on, his antennae began to vi- 
brate rapidly and continuously over her 
body. Within seconds he attempted to 
mount her. The female resisted by stridu- 
lating strongly, raising herself on her legs 
and flexing the apex of her metasoma 
slightly towards her coxae. As soon as the 
male began stroking her with his anten- 
nae, she became subdued, stopped strid- 
ulating and became absolutely still. Once 
on the female, the male continued to flick- 
er his antennae, continuously stroking her 



head and the anterior part of her meso- 
soma (Fig. 3). The female remained in a 
frozen position, with her antennae con- 
cealed under her deflexed head. After a 
period ranging from several seconds to a 
couple of minutes, depending on her re- 
action, the male gradually manoeuvred 
posteriorly on the female so that his gen- 
italia could be inserted into her genital 
opening. If she became restless the strok- 
ing of her body by his antennae intensi- 
fied. If she became more restless, he 
would quickly resume his initial more an- 
terior position. 

After moving posteriorly, the male 
grasped the female laterodorsally at the 
midlength of the first metasomal segment 
with his mandibles; extruding his genita- 
lia he began prodding her genital opening 
with them. Often, while the male was 
prodding her genital opening, the female 
would wander around the petri dish with 
him still mounted on her back. If she be- 
came too agitated or began moving too 
quickly, the male withdrew his genitalia, 
disengaged his mandibles and again be- 
gan stroking her with his antennae. As 
soon as actual genital union occurred, the 
female became motionless, thrusting her 
body forward, tucking her forelegs under 
her head, with the middle and hind legs 
placed laterally and supporting her. Her 
entire body was more or less straight with 
the metasoma lifted and the head against 
the substrate. Her ovipositor was extrud- 
ed (Fig. 4), a condition which may be nec- 
essary for successful copulation in mutil- 
lids since it has been observed in other 
species (Brothers 1972, Bayliss and Broth- 
ers 1996). Genital union lasted between 
60-100 seconds, during which time the 
male continuously stroked the female 
with his legs and antennae. Throughout 
genital union, the parameres remained 
outside the body of the female, lateral to 
her genital opening, while the rest of the 
male's genitalia extended into the female. 

Immediately following separation of the 
genitalia, the male, poised posteriorly on 



Journal of Hymenoptera Research 







Figs. 3-6. Dolichoiiiiitilhi si/a'n7.v, mating behaviour, diagrammatic. 3, mounted 6 stroking 9 with antennae. 
4, posture immediately before copulation. 5, posture immediately after copulation, b, posture several seconds 
after copulatit>n. 



the metasoma of the female, extended and 
straightened his metasoma, thrusting the 
tip high into the air, and retracted his gen- 
itaha (Fig. 5). Suddenly, without warning, 
the male rushed forward over the female, 
coming to rest on her mesosoma. He 
dropped his head on to hers and lifted the 
posterior part of his metasoma high into 
the air, almost perpendicular to the sub- 



strate, while swaying back and forth for 
approximately 5-10 seconds before dis- 
mounting (Fig. 6). Inimediately upon gen- 
ital separation the female retracted her 
ovipositor and bent her metasoma for- 
ward between her legs. She bent her head 
down and nibbled the metasomal tip with 
her mouthparts. This behaviour has pre- 
viously been reported as unique for the 



Volume 10, Number 1, 2001 



Formicidae within the Hymenoptera (Wil- 
son 1962, Parish 1972), but has never been 
recorded following copulation. Its func- 
tion is not obvious but, despite the fact 
that no extruding material could be seen, 
it is possible that part of the material de- 
posited by the male is a nuptial donation 
which the female consumes and uses as 
food or as some chemical signal. The fe- 
male did not extrude and withdraw her 
sting as observed by Brothers (1972) in 
Pseudomethoca frigida, and by Bayliss and 
Brothers (1996) in Tricholabiodes spp. After 
dismounting, the male usually began 
grooming himself thoroughly. It was sev- 
eral seconds before the female began to 
wander around the petri dish again. 

In subsequent encounters immediately 
following mating, the male's response to 
the female was one of apparent hostility; 
he rushed at her, fluttering his wings and 
bumping into her from behind. After sev- 
eral seconds of such treatment, and with 
no possible escape from the petri dish, the 
female stopped moving and hunched up, 
curling her head and antennae under her 
body and tucking her legs against her 
sides. The male continued to bump her 
from behind, rushing at her with wings 
flapping, apparently attempting to drive 
her away. Subsequent encounters between 
the male and female were of shorter du- 
ration, with continued aggressive behav- 
iour exhibited by the male towards the fe- 
male. Similarly, if two previous recently 
mated adults were again placed together, 
the male immediately became aggressive 
towards the female, chasing her with 
wings fluttering and bumping her, almost 
pouncing on her. The male became more 
aggressive to the mated female the longer 
they were kept together. There was never 
an attempt by the male to mount an al- 
ready mated female. As previously ob- 
served in P. frigida (Brothers 1972), the at- 
tractiveness of a mated female mutillid ap- 
pears to diminish rapidly after mating. Af- 
ter several days the aggressive behaviour 



of the male towards the female had van- 
ished, with him totally ignoring her. 

Unlike the situation in some other Smi- 
cromyrmina (Mutillini), some Myrmosi- 
nae and the Rhopalomutillinae (Brothers 
1975, 1989), where the male often trans- 
ports the female in flight before settling 
and mating or may even mate in flight, in 
D. sycorax no attempt was made by the 
male to fly and carry the female, and mat- 
ing took place on the substrate in an up- 
right position. The absence of phoretic 
copulation is probably because the male is 
about the same size as the female or even 
smaller. 

Grooming 

There are no differences in cleaning 
techniques between the sexes (except for 
those involving the wings). If an individ- 
ual is extremely dirty it first partially 
cleans the posterior part of the body; oth- 
erwise grooming proceeds antero-posteri- 
orly. 

Head. — The antennae, which are the 
most frequently groomed structures, are 
cleaned using the antenna cleaners on the 
front legs, either by double-antenna scrap- 
ing (both antenna cleaners are simulta- 
neously passed distad along the respective 
ipsilateral antenna) or single-antenna 
scraping (one antenna at a time is 
groomed by the ipsilateral antenna clean- 
er; the different antennae are usually 
groomed consecutively). During double- 
antenna scraping, the head remains still 
with each antenna placed in its antenna 
cleaner and then drawn between the spur 
and basitarsus from base to apex three to 
four times by movement of the forelegs. 
During single-antenna cleaning, the leg is 
lifted over the antenna which is placed in 
and pulled through the antenna cleaner by 
tilting the head backwards and simulta- 
neously moving the leg away from the 
head. Sometimes, more often in the fe- 
male, there is simultaneous grooming of 
one antenna using the antenna cleaners of 
both ipsi- and contralateral forelegs. The 



Journal of Hymenoptera Research 



surface of the head is cleaned by both 
forelegs separately or simultaneously. If 
the head is cleaned by only one foreleg, it 
is tilted to one side and brushed postero- 
anteriorly with short rapid strokes. The 
brushing of the head is usually followed 
by single-antenna scraping. The foreleg 
calcaria are used for cleaning the mandi- 
bles, while both maxillary and labial palpi 
are cleaned similarly to double-antenna 
scraping, where the palpi are either singly 
or simultaneously pulled rapidly through 
the antenna cleaner of the ipsilateral fore- 
legs. While one foreleg is cleaning the ip- 
silateral antenna, the other might be clean- 
ing the palpi. 

Body. — Cleaning of the dorsal and lat- 
eral parts of the mesosoma was never ob- 
served. The anterior part of the mesoso- 
ma, including the neck region, is cleaned 
with the forelegs separately or simulta- 
neously. The mesosternum is cleaned by 
the calcar and basitarsus of the foreleg; the 
calcar is first angled away from the basi- 
tarsus, then pushed down the length of 
the mesosoma between the coxae, ending 
with the foreleg rubbing laterally against 
the ipsilateral middle leg. The dorsal and 
lateral parts of the metasoma are cleaned 
by both hind legs, using alternating or si- 
multaneous strokes. While grooming the 
metasoma, the wasp balances on its front 
two pairs of legs, with the entire body 
slightly arched and the wings folded dor- 
sally. Often only one side of the metasoma 
is cleaned, using the ipsilateral leg. The 
longer tibial spur, which is that mainly 
used, is angled away from the tibia. The 
metasoma is first cleaned proximally, then 
sequentially more distally using longer 
strokes each time, the first one or two seg- 
ments being cleaned before proceeding to 
the more distal segments. The sides are 
groomed first, followed by the dorsal sur- 
face and then the sterna which are cleaned 
by a single hind leg. While the middle and 
hind legs clean the metasoma, the animal 
balances on its head with the forelegs sup- 
porting it laterally but close to the head. 



Although the hind legs are predominantly 
responsible for cleaning the metasoma, the 
middle legs might assist by making sev- 
eral strokes down the sides. No concentra- 
tion of attention to grooming of the felt 
lines (laterally on the second metasomal 
tergum) or the metasomal apex was ob- 
served, although these areas may be 
sources of pheromones or other chemicals. 

Legs. — The legs are cleaned sequential- 
ly, anterior to posterior. The fore legs, if 
very dirty are first rubbed against each 
other. The entire foreleg is then rubbed 
against and pulled between the spur and 
basitarsus of the ipsilateral middle leg. 
The ipsilateral middle leg is not moved. 
Alternatively, the fore leg is positioned 
ventrally along the length of the body, and 
the spur and basitarsus of the ipsilateral 
middle leg is scraped down its entire 
length and then shaken. Cleaning of the 
forelegs, in particular the tarsi and apical 
portions of the tibiae, using the mouth- 
parts (otherwise known as foreleg nib- 
bling) was never observed, although Bas- 
ibuyuk and Quicke (1999) noted this as 
commonly occurring in Mutillidae. The 
middle legs are groomed separately, using 
the tibial spurs and basitarsi of both hind 
legs. The hind legs are individually 
cleaned by the spurs and basitarsi of the 
ipsilateral middle and contralateral hind 
legs. The hind leg remains still. A hind leg 
is first cleaned distally, then sequentially 
more proximally by cleaning a longer sec- 
tion each time that the other legs are 
rubbed against it. The hind leg used for 
cleaning, if very dirty, is shaken or the 
basitarsi of both hind legs are rubbed to- 
gether. 

Wings. — The left and right wings, like 
the antennae, are cleaned either separately 
or simultaneously, using the spur and bas- 
itarsus of the ipsilateral hind leg. This is 
not to be confused with ichneumonid-type 
wing grooming where both pairs of raised 
wings are groomed simultaneously be- 
tween the tibia and tarsus of each respec- 
tive ipsilateral hind leg (Behaviour 16 



Volume 10, Number 1, 2001 



(Basibuyuk and Quicke 1999)). Unlike ich- 
neumonid-type wing grooming, where the 
wings are cleaned while in a horizontal 
position (Basibuyuk and Quicke 1999), in 
D. sycorax the wings are orientated ventro- 
laterally to the metasoma. While the fore- 
wings are cleaned, the hind wings are po- 
sitioned laterally, perpendicular to the 
body and horizontal to the substrate. The 
wing, while being cleaned, always re- 
mains between the body and the hind leg. 
While the dorsal surface of the forewing 
is being cleaned, the costal margin is ori- 
entated ventrally with the dorsal surface 
facing outwards. The metatibia and me- 
tabasitarsus, remaining lateral to the wing, 
slowly comb it in a backward-downward 
motion. After 3-4 strokes the hind leg is 
cleaned. The posterior margin of the fore- 
wing is cleaned once the dorsal surface 
has been combed; it is gripped and pulled 
between the spur and basitarsus. There- 
after the forewing is orientated so that the 
costal margin is dorsally placed, with the 
ventral surface facing outward. The ven- 
tral surface of the forewing is cleaned sim- 
ilarly to the dorsal surface. The hind wing 
is cleaned in a sequence similar to the 
forewing. The latter returns to a horizontal 
position along the body after scraping, 
though at a greater distance from the body 
than in the normal resting position. The 
hind wings are similarly flipped as the 
forewings, depending on the surface being 
cleaned. When the wings of both sides are 
being cleaned simultaneously, the animal 
balances on its front and middle legs, and 
when cleaning the wings separately, the 
wasp shifts its weight to either the left or 
right legs, arching the mesosoma away 
and the metasoma towards the wing that 
is being cleaned. 

Compared with Tricholabiodes spp. (Bay- 
liss and Brothers 1996), D. sycorax displays 
relatively few differences in grooming 
techniques but does tend to exhibit a 
greater repertoire of grooming behav- 
iours. 



Host Relations 

Specimens of DolichomiitiUn have been 
reared from the mud nests of various host 
wasps (specially sphecids of the tribe Sce- 
liphrini) and appear to be restricted to 
such hosts. Nonveiller (1996) gave the host 
of D. giiineensis (Fabricius) (= D. similUma 
Bischoff) as an unidentified species of 
Chah/bion, and that of D. scuteUifera (An- 
dre) (= D. conigera (Andre)) as an uniden- 
tified species of Sceliphron. Krombein & 
Walkley (1962) recorded D. minor minor 
Bischoff as a parasitoid of Sceliphron spiri- 
fex; we have seen a female specimen of D. 
m. minor collected on a Sceliphron nest at 
Mkuzi Game Reserve, KwaZulu-Natal 
(27°37'S, 32°14'E) on 3-6.iii.1990 by A. 
Weaving (DJB collection) and a female of 
D. }n. minor reared from a mud nest of 
"Cyphononyx antennatus (Smith)" (Pompi- 
lidae) collected at Durban on 10. i. 1945 by 
Marley (South African Museum collec- 
tion). Weaving (1994a, 1994b, 1995) re- 
corded the hosts of D. heterodonta Bischoff 
in KwaZulu-Natal as Auplopus femoralis 
(Arnold) (Pompilidae), Tricarinodynerus 
guerinii (Saussure) and Afreumenes aethiop- 
icus (Saussure) (both Vespidae, Eumeni- 
nae), all species using mud in nest mak- 
ing. (The mutillid was probably misiden- 
tified, however, since that species only oc- 
curs further north.) 

Peringuey (1898) reared both sexes of D. 
sycorax from the mud nests of S. spirifex 
on several occasions. Skaife (1953:325) re- 
ferred to "D. guineeiisis" (actually D. sy- 
corax) as having been reared from S. spi- 
rifex, but gave no authority for this, and 
may have been referring to Peringuey's 
specimens. (Incidentally, Skaife's figure 
163, captioned as being of both sexes of D. 
giiineensis "Parasitic on solitary bees" (sic), 
shows a male which looks like a species 
of Stenomutilla and a female which is prob- 
ably a species of Dasylahroides.) Weaving 
(1994b, 1995) recorded S. spirifex, T. guer- 
inii and Synagris analis Saussure (Vespi- 
dae, Eumeninae) as hosts of D. sycorax, 



Journal of Hymenoptera Research 



and provided considerable information on 
the biologies of the hosts and the influence 
of nest type and construction on parasit- 
ism rates. In the Albany Museum (Gra- 
hamstown) there are three female and 
three male specimens of D. si/corax reared 
from a nest of S. spirifex which also yield- 
ed one female and one male of the host 
and was collected by NJ. Myers at the To- 
bacco Research Station, Trelawney, Zim- 
babwe, January /February 1954 (dets F.W. 
Gess). In addition to these records and the 
same host relationship recorded in this pa- 
per, D. sifcorax has also been reared from 
multicellular mud nests of a species of eu- 
menine vespid, possibly Delta maxillosa 
(de Geer) (det. C.F. Jacot-Guillarmod) or 
S. analis (see Weaving 1995), collected by 
DJB at Lake Sibaya, KwaZulu-Natal on 
13-25 March 1968. That nest yielded no 
host specimens, but produced six mutil- 
lids, five females (10-18 mm long) and one 
male (10 mm long). The considerable dif- 
ference in sizes of individuals in this 
clutch is notable. The largest female has 
golden brown pubescence replacing the 
black pubescence of the other specimens 
and thus appears very different in color- 
ation. (Bischoff (1920) described a similar 
female specimen of this species as form 
aurata, and this phenomenon was first not- 
ed by Andre (1899:35) for other mutiUids.) 
The host range of D. sycorax is thus greater 
than previously thought, although in all 
cases mud is used for nest construction, 
whether as free multicellular aerial nests 
or forming the nest closures and cell par- 
titions in cavity nests. 

Also interesting is the fact that for two 
of the three nests from which multiple 
mutillids emerged, a single male was pro- 
duced. This may indicate a tendency to- 
ward a biassed sex ratio as is found in 
some other parasitoid hymenopterons, 
specially those which develop gregarious- 
ly or quasi-gregariously, and which pro- 
duce a single male that emerges early and 
mates with his sisters as they emerge (e.g. 
see Hardy 1994). Additional evidence is 



obviously needed, but partial support 
may be derived from the observation that 
males of DolicJwmutilla are very much rar- 
er in collections than are females. 

ACKNOWLEDGEMENTS 

We wish to thank Ray Miller and Johan Kotze of 
the University of Natal (Pietermaritzburg) and the 
KwaZulu-Natal Nature Conservation Service for the 
specimens. Fred Gess of the Albany Museum kindly 
supplied information on specimens in his care. Fi- 
nancial assistance was provided by the Foundation 
for Research Development and the University of Na- 
tal Research Committee. 

LITERATURE CITED 

Andre, E. 1899-1903. Les MutiUides. Species 
d'Hyiuaiopteyes d'Euroye at d'Algerie 8:1-479, pi. I- 
XV.' 

Basibuyuk, H. H. & D. L. J. Quicke. 1999. Grooming 
behaviours in the Hymenoptera (Insecta): poten- 
tial phylogenetic significance. Zoological jounml of 
the Liuuemi Society 125: 349-382. 

Bayliss, P. S. & D. J. Brothers. 1996. Biology of Tri- 
cholalnodes Radoszkowski in southern Africa, 
with a new synonymy and review of recent bi- 
ological literature (Hymenoptera: Mutillidae). 
loHiital of Hymenoptera Researcli 5: 249-258. 

Bischoff, H. 1920-21. Monographic der Mutilliden Af- 
rikas. Archiv fur Naturgeschichte 86(A): 1-830, 1 
map. 

Brothers, D. J. 1972. Biology and immature stages of 
Pseudomethoca f. fiigida, with notes on other spe- 
cies (Hymenoptera: Mutillidae). University of 
Kansas Science Bulleti)i 50: 1-38. 

Brothers, D.]. 1975. Phylogeny and classification of 
the aculeate Hymenoptera, with special reference 
to Mutillidae. University of Kansas Science Bulletin 
50: 483-^48. 

Brothers, D. J. 1989. Alternative life-history styles of 
mutillid wasps (Insecta, Hymenoptera), pp. 279- 
291. /;; Bruton, M.N. ed.. Alternative Life-Histori/ 
Sti/les of Aniiiuils, Kluwer Academic Publishers, 
Dordrecht. 

Parish, D. J. 1972. The evolutionary implications of 
c|ualitative variation in the grociming behaviour 
of the Hymenoptera (Insecta). Animal Behaviour 
20: 662-676. 

Ferguson, W. E. 1962. Biological characteristics of the 
mutillid subgenus Photopsis Blake and their sys- 
tematic values. University of California Publications 
in Entomologi/ 27: 1-91. 

Gerstaecker, [A.] 1857. [Descriptions.] in: i'eters, 
Ubersicht der von ihm in Mossambique aufge- 
fundenen und von Hrn. Dr. Gerstiicker bearbei- 
teten Hymenopteren aus den Familien der Cra- 
bronites, Sphegidae, Pompilidac und Heterogy- 



Volume 10, Number 1, 2001 



na. Mo)intsbcrichtL'n Akadciuic dcr Wissciischaftcii 
Berlin 1857, pp. 509-513. 

Gerstaecker, A. 1862. Hymenoptera, Hautfliigler. In: 
Peters, Naturanssciifiiiiaftllchc Rclsc imcli Mossivn- 
biqiic aiif Bcfehl seiner Majestiii dcs Konigs Friedrich 
Wilhelm IV in den Jahren 1842 bis 1848 ausgefiihrt. 
Zoologie, V. Iiisecten und M\/riopedes. Reimer, Ber- 
lin, pp. 438-526. 

Hardy, I. C. W. 1994. Sex ratio and mating structure 
in the parasitoid Hymenoptera. Oikos 69: 3-20. 

Krombein, K. V. & L. M. Walkley. 1962. Three hy- 
menopterous parasites of an African mud-daub- 
er wasp, Sceliphrou spirifex (L.) (Hymenoptera). 
Proceedings of the Entomological Societi/ of Washing- 
ton 64:78. 

Nonveiller, G. 1996. Remarques sur Dolichoinutilla 
guineensis {¥., 1793) Dolichoniutilla si/corax (Smith, 
1855) et sur certains hotes du genre Dolichonm- 
tilla Ashmead, 1899 [Hymenoptera, Mutillidae]. 
Revue frangiise d'Entoniologie 18: 31-34. 

Peringuey, L. 1898. Description of some new or little 
known South African Mutillidae in the collection 
of the South African Museum. Annals of the South 
African Museum 1: 33-94. 



Skaife, S. H. 1953. African Insect Life. Longmans 
Green, London etc. 

Weaving, A. J. S. 1994a. Notes on nesting behaviour 
in two Afrotropical auplopine wasps, Auplopus 
vitripennis Smith and A. femoralis (Arnold) (Hy- 
menoptera: Pompilidae). The Entomologist 113: 
140-153. 

Weaving, A. J. S. 1994b. Nesting behaviour in three 
Afrotropical trap-nesting wasps, Chalybion laevi- 
gatum (Kohl) Proepipona meadewaldoi Bequaert 
and Tricnrinodyiwrus gueriiiii (Saussure), (Hyme- 
noptera: Sphecidae, Eumenidae). The E}itomolo- 
gist 113: 183-197. 

Weaving, A. J. S. 1995. A comparison of nesting suc- 
cess and nesting habits in some Afrotropical acu- 
leate wasps, with particular reference to nest par- 
asites (Hymenoptera: Sphecidae, Eumenidae). 
A)i)uils of the Cape Provincial Museums (Natural 
History) 19: 181-224. 

Wilson, E. O. 1962. Behavior of Daceton arnugerum 
(Latreille), with a classification of self-grooming 
movements in ants. Bulletin of the Museum of 
Comparative Zoology 127: 401-422. 



J. HYM. RES. 
Vol. 10(1), 2001, pp. 10-15 

The Number of Sex Alleles (CSD) in a Bee Population and its 
Practical Importance (Hymenoptera: Apidae) 

GiSLENE Almeida Carvalho 

Departamento de Genetica da Faculdade de Medicina de Ribeirao Preto da Universidade de 
Sao Paulo, Ribeirao Preto, Sao Paulo, Brazil 



Abstract.— This work was carried out to monitor the number of xo sex alleles (CSD — Comple- 
mentary Sex Determination) in a limited population of McUpona scutellaris Latreille (Apinae, Me- 
liponini) and to verify if introduction of inseminated queens from distant populations is a good 
and practical method to avoid extinction of small ones. Twenty-two colonies ot Melipona scutellaris 
were brought from Len^ois, Bahia (12°34'S; 41°23'W) to Uberlandia (18°52'56"S; 48°12'55"W), 
500km south of the southern edge of its geographical distribution. Thirty foreign queens were 
introduced from 1992 to 1995, and the number of xo sex alleles oscillated from 7.0 to 36.0. The 
number of sex alleles was studied using a formula modified from the one of Laidlaw et al. (1956) 
n = 2M(N + 1)/(H+1) (where n is the number of xo sex alleles in the population, N is the total 
number of colonies sampled of this population, H is the number of colonies that produced diploid 
males and M is the number of males that mated with the queen). These results showed that the 
introduction of inseminated queens serves to maintain the variability of sexual alleles and, con- 
sequently, allow the existence of small populations of Meliponini. These data are being taught to 
Melipoiia beekeepers to improve their bee yards and help in protecting the species from extinction. 



Brazilian stingless bees (Hymenoptera, 
Apidae) are among the main pollinators of 
the native Brazilian flora, varying from 
30% to 90% of the native plants, according 
to the ecosystem they inhabit (Kerr et al. 
1994). These bees belong to the subfamily 
Apinae, tribe Meliponini, and at least 
three species, Melipoiia scutellaris Latreille, 
M. compressipes fasciculata Smith and M. 
beecheii, were domesticated by native Pre- 
colombian populations, two in Brazil 
(Maranhao and Pernambuco States) and 
one in Mexico (Yucatan). Besides being 
good pollinators, two bees, the African 
honey bee and Uruq:u {Melipona scutellaris) 
are the main producers of honey in north- 
eastern Brazil. Melipona honey reaches a 
price 4 to 10 times higher than honey bees 
because of lower production by individual 
colonies and the preference of local pop- 
ulations. 

More than 350 species of Meliponini are 
distributed in the Tropical and Subtropical 



Zones of South and Central Americas, Ma- 
laysia, India, Indonesia, Africa and Aus- 
tralia. Unfortunately about 100 species of 
stingless social bees are being seriously 
threatened with extinction as a conse- 
quence of the current forest destruction. In 
Brazil, annual deforestation increased 20% 
in the last 3 years due to the arrival of 
Chinese and Malaysian lumber compa- 
nies. 

Besides the forest destruction that is ac- 
companied by both forest fires and plan- 
tations of grass, soybean, rubber trees, 
guarana, pepper, Brazil nuts and tropical 
fruits, there is the serious problem of CSD 
(Complementary Sex Determination) that 
in bee happens with one main gene (.yo- 
sex alleles). According to Mackensen 
(1951) sex in Apis mellifera is determinated 
by xo sexual alleles, where heterozygous 
larvae become females, hemizygous are 
males and homozygous are diploid males. 
This genetic system by xo-sex alleles de- 



Volume 10, Number 1, 2001 



11 



Table 1. Production of males in colonies of Melipoiw scutcllnris and estimates xo alleles using n = 2M(N 
+ 1)/(H + 1) where n = number of xo, N is the total number of hives, H is the number of hives that produced 
50'/ci diploid drones and M is the number of males that inseminated the queen. 











Number of 


Number of 












colonies that 


colonies that 






Total number of 






produced 30"^, 


produced 75";. 






colonies in 




Number of 


female: 50"., 


female :25''m 


Estimated 


Year of 


Miiipona apiary 


Number of 


colonies produced 


male (mated with 


male (mated 


number of 


sampling 


in each vear 


colonies sampled 


100",, ot female 


one male) 


with two males) 


vo CSD alleles 



1991 


50 


13 


<-) 


1 


3 


24.5 


1992 (a) 


80 


27 


19 


7 


1 


7.86 


1993 (b) 


72 


15 


12 


3 





8.00 


1994 (c) 


79 


16 


13 


1 


2 


28.34 


1995 (d) 


66 


8 


7 





1 


36.00 


1997 


65 


7 


5 


1 


1 


12.00 


1998 


70 


9 


7 


1 


1 


15.00 


1999 


60 


29 


23 


5 


1 


11.67 


Total 


— 


124 


95 


19 


10 


23.86 



(a) introduction of 13 queens of Piata, BA (14 June 1992) 

(b) introduction of 3 queens of Catu, BA (19 July 1993) 

(c) introduction of 11 queens of Len(;6is, BA (24 May 1994) 

(d) introduction of 3 queens of Len^ois, BA (26 March 1995) 



termination is named "Complementary 
Sex Determination" (CSD) and was first 
described in the parasitoid wasp Br aeon 
hebetor by Whiting (1943). He proposed 
that sex was regulated by a series of alleles 
segregating at a single locus. Camargo 

(1979) found the same CSD genetics for 
Melipotia quadrifnsciata, Kerr (1987) for M. 
compressipes fasciculata, and Carvalho et al. 
(1995) for M. scutellnris. 

Kerr (1987) and Kerr et al. (1988) pro- 
posed that the xo" gene of primitive pop- 
ulations is still found in endogamous pop- 
ulations of Hymenoptera. It is mutated 
rarely, but constantly, to xo\ xo-, . . . .vo-'^ 
which were selected in panmitic popula- 
tions giving origin to a series of multiple 
alleles that are involved in sex determi- 
nation (CSD). 

Yokoyama and Nei (1979) and Cornuet 

(1980) have shown that the number of 
CSD alleles maintained in a limited bee 
population depends directly on its size. 
Woyke (1980) demonstrated that in Apis 
mellifera the minimum number of CSD al- 
leles that allows a population to survive is 
six. Our experience with three Melipona 
species indicates that six is also the mini- 
mum number of xo alleles required to 



maintain restricted populations. Bellow 
six the population decreases rapidly due 
to the formation of diploid males. Stou- 
thamer et al. (1992) and Heimpel et al. 
(1999) also observed the same in some hy- 
menopteran parasitoids. 

Unlike Apis mellifera queens that mate 
with 17 to 15 males (respectively Adams 
et al. 1977, Lobo and Kerr 1993), queens of 
stingless bees (Meliponini) mate with one 
or two males, rarely more (Kerr 1969, 
Contel and Kerr 1976, Paxton et al. 1999). 
Our data with Melipona scutellaris showed 
about 8% of crosses occurred with 2 males 
(Table 1). In small populations of Melipona 
with 6 xo alleles, Vs of the new colonies 
will produce diploid males (Kerr and Ven- 
covsky 1982). Many known panmitic Hy- 
menoptera have A'O" sex alleles and have 
developed different methods to avoid the 
production of triploid females: a) in many 
Hymenoptera diploid males are semi-le- 
thal or almost sterile (Inaba 1939, Mac- 
Bride 1946, Hung et al. 1974, Naito and Su- 
zuki 1991, Stouthamer et al. 1992, El Agoze 
et al. 1994); b) in Apis mellifera the workers 
eat the diploid male larvae (Woyke, 1980); 
c) in Melipona soon after they emerge from 
the brood cells, workers kill both diploid 



12 



Journal of Hymenoptera Research 



males and the inseminated queen that is 
producing them (Camargo 1979, Kerr 
1987, Kerr et al. 1996); d) increasing the 
number of .ro-alleles and e) multiple mat- 
ing dimishes the genetic load effect of 2n 
males. 

Adams et al (1977) and Lobo and Kerr 
(1993) respectively estimated the number 
of Ao-alleles for open populations of Apvs 
mellifern at 18.9 and 15.7, and Kerr (1987) 
in Melipojia compressipes found it was 20.0. 
Using data based in the formula of Cor- 
nuet (1980), Kerr and Vencovsky (1982) es- 
timated that in order to maintain 6 .Yo-al- 
leles, the Melipona scutellaris population 
must contain 44 colonies or more. If the 
number of colonies is below 44 this whole 
mini-population is bound to be eliminated 
in a few generations. Carvalho et al. (1995) 
cited many examples of the extinction of 
small populations of stingless bees. This 
means that the decline of the number of 
xo sex alleles in stingless bees (Meliponini) 
is fatal and leads a population and even 
the species to elimination. Falk (1991) em- 
phasized the fact that there is a worrying 
decline in the Aculeate population in 
Great Britain, with nearly half of the spe- 
cies described considered to be under 
threat. Of course, there is a genetic load 
associate with CSD sex determination in 
bees as it was demonstrated by Kerr 
(1975) and Werren (1993). 

The objective of the research presented 
in this paper was to monitor the number 
of xo alleles year by year in a limited pop- 
ulation of Melipona scutellaris and to study 
the effect of the introduction of inseminat- 
ed queens in this population. Descendants 
of 22 colonies that came from the forests 
near Len^ois (Bahia) were maintained in 
Uberlandia (18°52'56"S; 48°12'55"W) about 
500 km South of the southern border of its 
natural distribution, that is, there was no 
feral population of Melipona scutellaris 
within a radius of 500 kilometers of Uber- 
landia. 



MATERIALS AND METHODS 

The following material was used: 22 col- 
onies of Melipona scutellaris collected ran- 
domly in the forest that surrounds Lengois 
(12°34"S; 4r23"W), Chapada Diamantina, 
Bahia, Brazil — 14 hives in 1988 and 8 hives 
in 1990. These colonies were divided and 
from their descendents 124 were sampled 
to be monitored. In order to count the 
number of .vo-alleles the technique of Kerr 
(1987) was used, that is: take one or two 
brood combs with young bees of a given 
colony and put it in the place of the moth- 
er colony to receive the adult bees. In 1 to 
10 days, one or several virgin queens 
emerge; four days after emerging one 
makes the nuptial flight and is inseminat- 
ed. After 5 to 15 days the new queen be- 
gins egg laying. I marked this queen on 
her thorax. When the oldest brood comb 
of this new queen contains pupae, a small 
piece of it is taken (with about 10 to 30 
pupae) and the number of workers, 
queens and males is counted. If the pro- 
portion fits 1:1 females: males, it indicates 
50% production of diploid drones and is 
indication of insemination by one male 
only. Diploid males are confirmed by cy- 
tological analysis (diploid males have 18 
chromosomes). If 25% males were pro- 
duced it indicated that two males had in- 
seminate this queen and if it was 12.5% 
three matings were indicated. Then the 
data are analyzed using the Laidlaw et al. 
(1956) formula n = 2M(N-I-1)/(H+1) that 
is better than n = 2MN/H; in both ec^ua- 
tions n is the number of xo sex alleles in 
the population, N is the total number of 
colonies sampled in this population, H is 
the total number of colonies sampled that 
produced diploid males and M is the 
number of males that inseminated the 
queen. This formula assumes that the xo 
alleles have equal frequency, that there is 
random mating, and there is less bias in 
populations smaller than 10. Fisher's test 
was used to analyze the variation of xo al- 
leles frequency by year. In order to mea- 



Volume 10, Number 1, 2001 



13 



sure the effect of the introduction of ge- 
netic material from outside the Uberlandia 
population as a search for a method of 
controlling the appearance of diploid 
males in small apiaries, 30 introductions 
of inseminated queens in orphaned colo- 
nies were carried out and these were: 13 
queens of Piata (Bahia) on 14 May 1992, 3 
queens of Catu (Bahia) on 19 July 1993, 11 
queens of Len^ois (Bahia) on 24 May 1994 
and 3 queens of Len^ois (Bahia) on 26 
March 1995. The mortality of egg and lar- 
vae of diploid males is near zero. 

RESULTS 

One hundred and twenty four samples 
of the new colonies made were taken from 
the Meliponn apiary of which 95 had 100% 
females in the first series of eggs laid by 
the new queen, 19 presented about 50% 
diploid drones and 10 about 25%; none 
produced 12.5%. The results of these sam- 
ples taken during 8 years are in Table 1. 

This data demonstrated that 65.5% of 
mating occurred with one male and 34.5% 
with 2 males. Then, the number of xo al- 
leles in this population is 8.18 (when 
queen mates with one male that has the 
same allele as she does) and 15.68 (when 
two males — one with the same allele and 
another with different allele) totaling 23.86 
xo alleles. 

Table 1 demonstrates the variation of xo 
alleles in this population as a consequence 
of sampling and matings with drones pro- 
duced in this same population. As these 
colonies were divided (in order to increase 
our population) since their introduction 
(1990), the probability that one queen 
mates with a drone that has the same al- 
lele is high because the population is 
small. The population in Uberlandia was 
formed of colonies from a native popula- 
tion in Bahia, and the number of alleles in 
that original population was expected to 
be about 20 (Kerr and Vencovsky 1982). In 
our 67 colonies (average per year) the av- 
erage number of alleles was 16.67. Look- 
ing at the data of 1991 to 1999, the number 



of sex alleles does not differ statistically 
between these years (Fisher's Test, 
P>0,05), which indicates the value of our 
method to avoid diminution of these al- 
leles. 

DISCUSSION 

The behavior of stingless bee workers 
killing both diploid males and the queen 
is how they control the appearance of 2n 
males. It causes many beekeepers and bee 
scientists who collect and maintain less 
than 44 hives per species to lose all or al- 
most all in short periods of time. Four ex- 
amples are: a) W. E. Kerr collected 14 
hives, in Parnaiba, SP, Brazil, of Melipoua 
marginatn in 1945. Six years later all had 
died; b) He collected 12 hives, in 1944, of 
Melipona qundrifascintn (also in Parnaiba, 
SP). In 1955 all had died; c) Eng. Agr. Ro- 
gerio M. O. Alves collected 11 colonies of 
M. qundrifascintn near Catu, Bahia, Brazil, 
in May 1990; by December 1991 he had 
eight colonies, and in June 1993 only one 
was left; d) Mr. Alvino Pianzolli collected 
20 colonies of uru^u-preto (Meliponn cnpix- 
nbn) in Domingos Martins, Espirito Santo, 
Brazil in 1973. In June 1993, he still had 8, 
because the forest is not very far from the 
meliponary, so that his queens can mate 
with males from the forest. The same case 
was observed by Nascimento et al. (1996) 
at Archipelago of Fernando de Noronha 
(Pernambuco, Brazil) for three species of 
Melipo)in. They found, in 1996, only 18 col- 
onies of M. compressipes out of 32 intro- 
duced, 11 of Meliponn suhnitida in 10 and 
none of M. scutellnris in 30 introduced in 
1982 by Kerr and Cabeda (1985). These ex- 
amples confirm the observations of Yo- 
koyama and Nei (1979) about the extinc- 
tion of small populations, and suggest the 
necessity of methods to avoid the decline 
of variability of sexual alleles. 

The introduction of 13 foreign queens in 
1992 and 3 in 1993 did not show its effect 
in the same year — the data show a decline 
in the number of alleles in 1992 and 1993. 
Due to the winter that followed the intro- 



14 



Journal of Hymenoptera Research 



auctions of 1992 and 1993, the colonies 
usually became weaker than in the rest of 
the year and there was no regular pro- 
duction of males. Therefore, these intro- 
duced queens will be genetically active 
through their males only for the next 
queens naturally superseded. 

When the queens were introduced in 
June or July (winter in the Southern Hemi- 
sphere) there were few or no males, but 
when they were introduced in May or 
March, the last season in which males are 
being produced, an apparent increase of 
the number of xo sex alleles was detected, 
reaching a value of n = 17.92, which is 
close to the values obtained by Kerr (1987) 
and Carvalho et al. (1995) for two Melipona 
species. Looking at the results of 1997 to 
1999, it appears to indicate a small decline 
in the number of xo alleles. As the statis- 
tical analysis (Fisher's Test) showed, there 
was no divergence among the number of 
alleles by year, indicating that the intro- 
duction of these queens maintained the 
genetic variability in the population in 
Uberlandia, thus avoiding the appearance 
of diploid males and the extinction of their 
population. 

Our data showed that the introduction 
of 3 to 4 inseminated queens per year in 
an apiary with about 65 colonies im- 
proved the genetic variability of the xo al- 
leles becoming a good alternative to avoid 
the production of diploid males. It can 
save the production of hundreds of bee- 
keepers who maintain small meliponaries 
because the production and sale of insem- 
inated queens is much easier than selling 
a complete bee colony. 

The main conclusion is: the method of 
introduction of 3 to 4 inseminated queens 
by year is good to avoid low sex allele di- 
versity and consequence production of 
diploid males (the Yokoyama and Nei ef- 
fect); it maintains the variability of xo sex- 
ual alleles and consequently allows the ex- 
istence of restricted populations of Meli- 
ponini ex-situ, that is outside their area of 
geographic distribution. 



ACKNOWLEDGMENTS 

1 thank Prof. Warwick Estevam Kerr tor the advices 
provided during this seven years work; Prof. Gismar 
Silva Vieira for helping in statistical analysis, Prof. 
Rogerio Marcos de Oliveira Alves for providing 
queens from Catu, BA; Mr. Jose Carlos Moraes for 
queens from Len^ois, BA. I thank Ms. Vania Alves 
Nascimento for her help in the cytological examina- 
tions. My acknowlegments are also to an anonymous 
referee that made important suggestions. CNPq (Con- 
selho Nacional de Pesquisa e Desenvolvimento Tec- 
nologico), FAPEMIG (Funda^ao de Amparo a Pes- 
quisa de Minas Gerais) and Bank of Brazil Founda- 
tion provided the financial support. 

LITERATURE CITED 

Adams, J., E. D. Rothman, W. E. Kerr, and Z. L. Pau- 
lino. 1Q77. Estimation of the number of sex alleles 
and queen matings from diploid male frequen- 
cies in a population of Apis iiicUifcrn. Genetics 86: 
583-596. 

Camargo, C. A. 1979. Sex determination in bees. Pro- 
duction of diploid males and sex determination 
in Melipona qundrifasciata. Journal of Apiciiltiiral 
Research 18 (2): 77-84. 

Carvalho, G. A., W. E. Kerr, and V. A. Nascimento. 
1995. Sex determination in bees. XXXIII. Decrease 
of xo heteroalleles in a finite population of Meli- 
pona scutellaris (Apidae, Meliponini). Brazilian 
journal of Genetics 18 (1): 13-16. 

Contel, E. P. B. and W. E. Kerr. 1976. Origin of males 
in Melipona subnitida estimated from data of an 
isozymic polymorphic system. Genetica 46 (3): 
271-277. 

Cornuet, J. M. 1980. Rapid estimation of the number 
of sex alleles in panmitic honeybee populations. 
Journal of Apicultural Research 19: 3-5. 

El Agoze, M., J. M. Drezen, S. Renaulst, and G. Per- 
iquet. 1994. Analysis of the reproductive poten- 
tial of diploid males in the wasp Diadrouiiis pul- 
clielliis (Hymenoptera: Ichneumonidae). Bulletiiii 
of Entomological Research 84: 213-218. 

Falk, S. 1991. A review of the scarce and threatened, 
wasps and ants of Great Britain. Peterborough, 
UK; Nature Conservancy Council for England. 
Research and Siirvei/ in Nature Conservation N"35, 
ii + 344pp. {Apicultural Abstracts 1992, vol. 43 (3): 
196-197). 

Heimpel, G. E., M. F. Antolin, and M. R. Strand. 1999. 
Diversity of Sex-determining alleles in Bincon lie- 
betor. Heredity 82: 282-291. 

Holloway, A. K., G. E. Heimpel, M. R. Strand, and 
M. F. Antolin. 1999. Survival of Diploid Males in 
Bracon sp near hebetor (Hymenoptera: Braconi- 
dae). Annals of the Entomologcial Societif of America 
92 (1): 110-116. 
Hung, A. C. F., S. B. Vinson, and J. W. Summerlin. 



Volume 10, Number 1, 2001 



15 



1974. Male sterility in the red imported tire ant, 
Solcnopsis liivicfti. Ammls of the Entoiiiolo'^cial So- 
ciet}/ of America 67: 909-912. 

Inaba, F. 1939. Diploid males and triploid females of 
the parasitic wasp Habrobacon pcctiiiophorae Wa- 
tanabe. Cytolo^in 9: 517-534. 

Kerr, W. E. .1969. Some aspects of the evolution of 
social bees (Apidae). Evoluiionary Biology 3 (4): 
119-175. 

Kerr, W. E. .1975. Population genetics studies in bees 
(Apide, Hymenoptera) I. Genetic load. Aiiais da 
Acadciiiia Brasilcira dc Ciciiicas 47 (2):319-334. 

Kerr, W. E. .1987. Biologia, manejo e genetica de Mc- 
lipona co))iprc$sipes fasciculafa Smith (Hymenop- 
tera: Apidae). Tese dc Professor Titular. UFMA. 
Sao Luis (MA), 141 pp. 

Kerr, W. E. and M. Cabeda. 1985. Introdui^ao de abel- 
has no territorio de Fernando de Noronha. Cicii- 
cia c Cult lira 37 (3): 467-471. 

Kerr, W. E. and R. Vencovsky. 1982. Melhoramento 
genetico em abelhas. I. Efeito do niimero de co- 
lonias sobre o melhoramento. Brazilian journal of 
Genetics 5: 279-285. 

Kerr, W. E., S. G. Monteiro, and H. A. S. Kerr. 1988. 
Sex determination in bees. XXV. Adaptative val- 
ue of the xo gene in its origin. Brazilia)! Jounml of 
Genetics 11 (2): 469 473. 

Kerr, W. E., V. A. Nascimento, and G. A. Carvalho. 
1994. Fia Salva(jao para os Melipom'neos? Anais 
do 2". Eiuwitro Sobre Abelhas, de Ribeirdo Preto 1: 
60-65. 

Kerr, W. E., G. A. Carvalho and V. A. Nascimento. 
1996. Abclha Urui^u: Biologia, Manejo e Coiiscrvaglo. 
Ed. Funda(;:ao Acangau, Belo Florizonte, MG. 144 
pp. 

Laidlaw, H. H., F. P. Gomes, and W. E. Kerr. 1956. 
Estimation of the number of lethal alleles in a 
panmitic population of Apis mellifera. Genetics 41 
(2): 179-188. 

Lobo, J. A. and W. E. Kerr. 1993. Estimation of the 
number of matings in Apis mellifera, extensions 



of the model and comparison of different esti- 
mates. Ethology, Ecology and Evolution 5: 337-345. 

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diploid males to penetrate the eggs. Genetics 31: 
224. 

Mackensen, O. 1951. Viability and Sex determination 
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509. 

Naito T. and H. Suzuki. 1991. Sex determination in 
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occurrence of triploid males, journal of Heredity 
82: 101-104. 

Nascimento, V. A., G. A. Carvalho, L. M. Cavalcante, 
and W. E. Kerr. 1996. Introdu(jao de abelhas no 
arquipelago de Fernando de Noronha. 4. A po- 
pulacjao de abelhas apos uma decada. Anais do U 
Encontro sobre abelhas de Ribeiriio Preto 2: 209-216. 

Paxton, R., N. Weibschuh, W. Engels, K. Hartfelder, 
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143-146. 

Stouthamer, R., R. F. Luck, and J. H. Werren. 1992. 
Genetics of sex determinaticin and the improve- 
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viromental. Entomology 21: 427-435. 

Werren, J. H. 1993. The evolution of inbreeding in 
haplodiploid organisms. /;;: Thornhill, N. W. 
(ed.). The Natural History of Inbreeding and Out- 
breeding, p. 42-59, University of Chicago Press, 
Chicago, IL. 

Whiting, P. W. 1943. Multiple alleles in complemen- 
tary sex determination of Habrobacon. Genetics 28: 
365-382. 

Woyke, J. 1980. Effect of sex allele homo-heterozigoz- 
ity on honeybee colony populations and their 
honey production. I. Favorable development 
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Apicultural Research 19 (1): 51-63. 

Yokoyama, S. and M. Nei. 1979. Population dynamics 
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626. 



J. HYM. RES. 
Vol. 10(1), 2001, pp. 16-28 



Distribution and Ethology of Priscomasaris Gess 
(Hymenoptera: Vespidae: Masarinae: Priscomasarina) 

in Namibia 

Sarah K. Gess 
Albany Museum, Grahamstown, 6139 South Africa 



Abstract. — Knowledge of the distribution and flower associations of the monospecific genus 
Priscomasaris is expanded. The nature of the provision, nesting situation, nest structure, and meth- 
od of nest construction are recorded for the first time. Prisavuasaris naiiiihiensis Gess has been 
encountered solely in the Mopane Savanna /Northern Namib transition and Dwarf Shrub Savanna 
(of Giess 1971), all records falling within the northwestern extension of the Nama Karoo Biome 
(of Rutherford and Westfall 1986, as adapted by Lovegrove 1993). Flowers visited for nectar and 
pollen are of the families Aizoaceae (non-Mesembryanthema) and Molluginaceae. Nesting aggre- 
gations were located in sparsely vegetated areas of horizontally presented, stabilized, sandy soil. 
The nest is a multicellular burrow with its entrance surmounted by a mud-turret and with each 
excavated cell containing a constructed mud-cell. Water is used in excavation and construction. 
Evidence for bivoltinism is presented. 



F.W. Gess (1998) described Priscownsaris 
uamibioisis Gess, a new genus and species 
of Masarinae (Hymenoptera: Vespidae) 
from Namibia. He discussed its position 
within the subfamily, placing it in a new 
subtribe Priscomasarina, a sister group of 
Paragiina and Masarina combined. Its 
ethology is of particular interest as it rep- 
resents the most primitive extant member 
of the Masarinae. F.W. Gess recorded fe- 
male water collection behaviour and two 
forage plants but, as nests had not yet 
been discovered, could make no comment 
on nesting behaviour other than to sug- 
gest that water is most probably used in 
nest construction. 

In March 2000 two nesting aggregations 
of P. namihiensis were discovered and in- 
vestigated by the present author. This in- 
vestigation forms the subject of the pre- 
sent paper. 

Distribution. —Previously published 
(F.W. Gess 1998) and new collection re- 
cords of P. namihiensis are listed in Appen- 
dix 1 under the vegetation types of Giess 
(1971) (Fig. 1) and biomes of Rutherford 



and Westfall (1986) as adapted by Love- 
grove (1993) (Fig. 2). It appears that P. na- 
mihiensis is a northern Nama Karoo spe- 
cies. 

Nesting areas aiid sites.— Two nesting ar- 
eas of P. naniibiensis were located, both in 
Dwarf Savanna Shrub to the east of the 
Mariental-Keetmanshoop road and rail- 
way track: one 5 km south of Mariental 
(Fig. 3) and the other 7 km south of Gib- 
eon railway siding. Both areas are associ- 
ated with drainage lines that form part of 
the Fish River catchment. Rainwater 
pools, resulting from thundershowers, 
were present in association with the rail- 
way embankment through which the 
drainage channels have passage (Fig. 4). 

The nests were aggregated above nor- 
mal flood level within sparsely vegetated 
sites. In the Gibeon nesting area two small 
aggregations were discovered, each of no 
more than 10 nests, 9 to 11 metres from 
the water source. The soil was compacted 
sand with an inclusion of small pebbles 
and with a sufficient clay element to make 
it maleable when mixed with water. The 



Volume 10, Number 1, 2001 



17 




Figs. 1-2. Map of Namibia showing the distribution, based on collection records (Appendix 1), of Pnsco- 
nwsnris un))iibieiisis (dots). 1, The vegetation types of Giess (1971): 1 = Northern Namib; 2 = Central Namib; 
3 = Southern Namib; 4 = Semi-desert and Savanna Transition (Escarpment Zone); 5 = Mopane Savanna; 6 
= Mountain Savanna and Karstveld; 7 = Thornbush Savanna; 8 = Highland Savanna; 9 = Dwarf Shrub 
Savanna; 10 = Saline Desert with Dwarf Shrub Savanna Fringe; 11 = Tree Savanna and Woodland; 12 = 
Camelthorn Savanna (Central Kalahari); 13 = Mixed Tree and Shrub Savanna (Southern Kalahari). 2, The 
biomes of Rutherford and Westfall (1986) as adapted by Lovegrove (1993) (S = Savanna; N = Namib Desert; 
SK = Succulent Karoo; NK = Nama Karoo). 



gravelly surface was scattered with peb- 
bles and small plants. Individual nests 
were sited next to a pebble or small plant 
(Figs. 5-10). 

In the Mariental nesting area numerous 
nests were distributed over a site c 100 
metres square. The soil was compacted 
sand of an even consistency with a suffi- 
cient clay element to make it maleable 
when mixed with water. The surface was 
compacted sand scattered with an occa- 
sional pebble and "mat" plant. Most com- 
monly nests were grouped beneath or be- 
tween the spreading branches of semi- 
prostrate Gisekia cifricana (Lour.) Kuntze 
(Molluginaceae) (Fig. 16), a forage plant of 
Priscomasaris, and of a "mat" forming spe- 
cies of buiigofera (Papilionaceae), the flow- 



ers of which it does not visit (Fig. 17). Less 
commonly nests were exposed but were 
then positioned next to a pebble or small 
plant. 

Water collection. — As stated by F.W. 
Gess (1998) females visit pools of water in 
drainage charmels and river beds in order 
to obtain water, apparently for use in nest 
construction. When they were common, 
very large numbers were present on and 
flying over the water. When filling their 
crops they always alight on the water sur- 
face, never at the water's edge. Whilst on 
the water surface the wings are held erect. 

Flower visiting. — Both sexes of P. nami- 
biensis visit flowers. Those recorded were 
all small shallow flow ers of Molluginaceae 
and Aizoaceae. The Molluginaceae were 



18 



Journal of Hvmenoptera Research 





Figs. 3-4. Net,ling area soutli ol Mariontal (24.40S 17.57E). 3, Habitat. 4, Water source. 



Volume 10, Number 1, 2001 



19 



pink flowered Gisekia africann (Lour.) 
Kuntze (Fig. 16) (between Palm and Kho- 
rixas) and white flowered Limeum of three 
species: L. nrgute-carinnUim Wawra & Peyr. 
(Fig. 12) (west of Khorixas, between Kho- 
rixas and Uis, and south of Mariental), L. 
myosotis H. Walter (Fig. 13) (between Kho- 
rixas and Uis), and L. sulcatum (Klotsch) 
Hutch. (Fig. 11) (southwest of Bullsport). 
The Aizoaceae (non-Mesembryanthema) 
were purplish-pink flowered Sesuvium se- 
suvioides (Fenzl) Verde. (Fig. 15) (south of 
Mariental) and pink and white flowered 
TriantJwmn parvifolia E. Mey. ex Sond. (Fig. 
14) (also south of Mariental). All yield nec- 
tar, easily imbibed by this short 
"tongued" wasp, and pollen, requiring no 
specialized "harvesting" behaviour. Other 
plants in flower at the foraging sites were 
not visited. 

Priscomasaris is not the sole visitor to the 
flowers of any of its forage plants. The 
Molluginaceae are widespread and are 
visited by a range of wasps and bees (Gess 
and Gess unpublished catalogue of flow- 
ers visited by aculeate wasps and bees in 
semi-arid areas in southern Africa). The 
Aizoaceae attract a narrow range of wasps 
and bees — S. sesuvioides is principally vis- 
ited by Ceramius damarinus Turner (Ves- 
pidae: Masarinae), which uses it as its 
source of provision for its nests (S.K. Gess 
1999), and a species of Parafidelia (Fideli- 
idae). T. parvifolia is principally visited by 
Quartiniella turneri Schulthess and several 
Quartinia species. 

At the site near Bullsport, where Pris- 
comasaris males and females were visiting 
the flowers of L. sulcatum, some males and 
an occasional female were collecting nec- 
tar from the extrafloral nectaries of young 
plants of the herb Chamaesyce glandulifera 
(Pax) Koutnik (Euphorbiaceae). 

Provision. — The cell provision consists of 
a compact, firm, roughly cylindrical mass 
of pollen and nectar, rounded at the ends 
and with undulations along its length in- 
dicating deposition of individual "loads" 
of the pollen and nectar mixture. The pro- 



vision mass, a "pollen loaf", remains loose 
within the cell. One complete "pollen 
loaf" was 8.17 mm in length and 3.0 mm 
in diameter (i.e. 0.12 mm less than the in- 
ner diameter of the constructed mud-cell). 
Samples of pollen from provision masses 
collected on 10, 22 and 31 March, and 2 
April were examined microscopically. 
Four distinct pollens of 17.5, 20, 25 and 30 
micromillimetres in diameter were pres- 
ent. These were compared with pollens 
from flowers visited by Priscomasaris and 
were found to match those of T. parvifolia, 
L. argute-carinatum, G. africana and S. se- 
suvioides respectively. The proportions of 
the different pollens varied. Samples taken 
on 22 March were mostly of G. africana 
and S. sesuvioides and those taken on 31 
March and 2 April were principally of T. 
parvifolia. 

Male behaviour. — No males were found 
in nests or at water. In the morning, before 
the appearance of the females, males were 
observed flying to and fro over nesting ag- 
gregations and visiting flowers. After the 
females had made their appearance males 
were observed mounting them, both on 
the ground in the vicinity of the nests and 
on flowers. It was not possible to observe 
copulation. 

Description and method of nest construc- 
tion. — The description of the nest of P. na- 
mibiensis is based on investigations of 62 
nests: four in the Gibeon nesting area on 
9 March; and 22, 13, 5 and 18 at the Mar- 
iental nesting area on 10, 22 and 31 March 
and 2 April respectively. The method of 
construction is based on investigations of 
nests together with observations on nest- 
ing behaviour. 

Description: The nest of P. namibiensis 
consists of a multicellular subterranean 
burrow (Figs. 19-23) surmounted by a 
curved, tubular mud-turret of the same di- 
ameter as the burrow opening and usually 
with its greater length parallel to the soil 
surface. The main shaft is vertical and is 
of equal diameter throughout its entire 
length. At its base it curves outwards to 



20 



Journal of Hymenoptera Research 




Figs. 5-10. Pn^cunKi^iin^ ii(Uiiihicii>i> w ith nest entrance turret. 5 and 6, Preparing to enter nest entrance turret. 
7, Entering nest entrance turret, ventral surface uppermost. 8, Preparing to leave nest to discard the mud- 
pellet held between mandibles. 9, Flying away from nest with a mud-pellet which will be dropped in the 
pellet-dropping area. 10, Adding mud to the rim of entrance turret. Actual length of wasp c 8 mm. 



Volume 10, Number 1, 2001 



21 




Figs. 11-16. Forage plants of Priscoiimris naniibiciisis. 11, Limcian sulcatuiii (Mollugmaccvtc), llowcis white, P. 
uamibiensis leaving after foraging on the flowers. 12, Limciim argutc-carinatutu (Molluginaceae), flowers white. 
13, Liuiciiiu inifosotis (Molluginaceae), flowers white. 14, Triantheuia pnrvifolia (Aizoaceae: non-Mesembryan- 
thema), flowers white or pink. 15, Scsuviiirii sesiwioidcs (Aizoaceae: non-Mesembryanthema), flowers purplish- 
pink. 16, Gisckia africmia (Molluginaceae), flowers pink; entrance turret of P. luviiibicn^h on left. 



22 



Journal of Hymenoptera Research 



form a short lateral shaft which terminates 
in a sub-horizontal excavated cell in which 
is a constructed mud-cell. A second cell 
terminates a similar lateral shaft which 
leaves the vertical shaft at the same depth 
but at an acute angle from the first such 
that the two cells lie close together. Fur- 
ther cells in similar pairs are positioned 
almost immediately below the first pair, 
each pair deeper than that preceding it, so 
that the cells form a "stack" to one side of 
the main shaft. The largest number of cells 
found was 13. 

Method of construction: When excavat- 
ing a burrow, P. namibiensis, using its 
mandibles, extracts soil as moist mud-pel- 
lets. As nest constructing females make 
frequent visits to water it is almost certain 
that regurgitated water is used for moist- 
ening the soil. 

At an early stage in shaft excavation the 
entrance turret is constructed from pellets 
extracted from the burrow. These are laid 
down around the rim of the entrance to 
the burrow in such a way that its inner 
diameter equals that of the main shaft (i.e. 
3 mm). At the outset of turret construction 
the thickness of the wall is c 1 mm but 
after the turret has reached a height of a 
few millimetres is reduced to 0.5 mm. Af- 
ter the turret reaches a height of c 6 mm, 
pellets are usually added in such a way 
that the turret curves over, typically until 
the opening is vertical, after which they 
are added evenly around the circumfer- 
ence of the turret and the resultant hori- 
zontal tube is extended parallel with the 
soil surface for a further 13-17 mm, sep- 
arated from it by a c 2 mm gap. 

A female, when entering a nest, alights 
on the upper surface of the turret facing 
towards the opening (Fig. 5) and then 
curves over its lip (Fig. 6), entering and 
progressing along the passageway ventral 
surface uppermost (Fig. 7). When leaving 
the nest during shaft excavation she re- 
verses the procedure emerging posterior 
end first, ventral surface uppermost and 
climbing out onto the upper surface (Fig. 



8). Thus before flying away she is facing 
towards the turret opening. Whilst build- 
ing the turret she remains curved around 
the rim (Fig. 10), rotating whilst placing 
mud with her mandibles and apparantly 
supporting and tamping it with the tip of 
her abdomen. 

When discarding a mud pellet, a female, 
holding the pellet in her mandibles, flies 
(Fig. 9) on a roughly circular path, drop- 
ping the pellets 10 to 30 cm from the nest. 
Usually the variation in distance of an in- 
dividual's flight path is not more than 7 
cm so that the pellets accumulate in a 
small area (Fig. 17). 

The walls of the shaft are stabilized and 
smoothed with the addition of water. The 
diameter of the shaft (3 mm) is maintained 
constant throughout its length. There is no 
turning "bulb" such as that found in the 
nests of most species of Cernmius (S.K. 
Gess 1996). At a depth of 60-80 mm (av- 
erage 69, n = 17) the shaft curves and after 
c 5 mm is expanded in the excavation of 
a cell of length c 15 mm and diameter c 5 



mm. 



Within the excavated cell a mud-cell 
(Figs. 19-23) is constructed, fitting closely 
within it but easily removed from it. On 
the outer surface evidence of deposition of 
mud-pellets is visible and a faint "fish 
scale" pattern similar to the more marked 
pattern exhibited by constructed cells of 
Celonites (S.K. Gess 1996) and Pseudomn- 
saris (Torchio 1970) is discernable (Fig. 21). 
The inner surface of the mud-cell is 
smoothed (Fig. 20). The closed inner end 
is rounded whereas the outside tip of the 
mud-cell is consistently markedly papil- 
late (Figs. 20-22). Apart from the tip, mud- 
cells are constant in diameter along their 
length, cigar-shaped, not ovoid. 

The source of soil for the construction 
of the mud-cells was not determined. No 
quarry site within the nest was found. In 
nests with newly constructed open mud- 
cells there did not appear to be an exca- 
vated or partially excavated cell which 
could have been the source. As there is no 



Vol I'M! II), Ni'MtUK !, 2(101 



23 







Fig. 17. Hntrance turret oi Pnscouiasans iiaiiiibicnsis ein right (arrow), discarded iniid-peliets on left (arrow), 
amongst the branches of a species of bidigofcm (Papilionaceae). 



turning "bulb" such a source is also elim- 
inated. The builders were carefully ob- 
served and were not bringing in soil from 
outside the nest. On numerous occasions, 
during nest investigation, loose dry soil 
was found in the main shaft from a depth 
of 50 mm downwards. It is highly unlikely 
that this soil had fallen into the shaft. It 
seems possible that, when a cell is exca- 
vated, extracted soil is not carried out as 
pellets but stored loose in the shaft and 
that the female coming and going from the 
nest can pass through the loose soil. This 
soil would then be available for mud-cell 
construction. 

The constructed mud-cell walls appear 
to be harder and more brittle than those 
of the turret. This suggests that either, as 
in drying of concrete, slower drying of the 
mud used in mud-cell construction results 
in a "stronger" cementing or that some- 
thing other than water is added to the 



mud mixture. As Celonites and Pseudomn- 
saris use nectar for bonding soil used in 
cell construction (S.K. Gess 1996 and Tor- 
chio 1970), nectar was considered a pos- 
siblity. One gram of cell wall was tested 
for sugar content. Total sugars extracted 
amounted to only 3 milligrams, 0.6 micro- 
grams of which were found to be glucose 
and 0.8 micrograms fructose. This appears 
to be too low a concentration of sugars to 
suggest deliberate use of nectar (Chris 
Whitely pers. com.) and therefore should 
be considered rather to be accidentally 
added from the crop which is used vari- 
ously to carry nectar and water. 

A mud-cell, in which oviposition and 
provisioning have been completed, is 
sealed with a mud-plug that fits into the 
neck of the cell, closing but not sealing it. 
After this plug has been constructed there 
is further addition of mud, extending 
across the plug and the rim of the mud- 



24 



Journal of Hymenoptera Research 



oH 



-o 




5cm 



Fig. 18. Plans of two nests of Priscoiniisinis luunibicii- 
sis. a and b, vertical in single plane, a -nest with first 
cell only, b — nest with 12 cells, c, transverse in single 
plane. 

cell, effectively sealing it. The remaining 
few millimetres of the lateral shaft are 
then filled with soil. Finally, where the 
shaft curves away from the vertical shaft, 
mud is used for sealing and plastering so 
that, when a nest with sealed cells is 
opened, there is no sign in the walls of the 
vertical shaft that any lateral shaft other 
than one leading to an open cell exists 
(Fig. 22). 

A first cell having been completed and 
sealed off, a second lateral shaft leading to 
a second cell is excavated (and a mud cell 
constructed within it) at the same depth 
as the first and in the same plane, but at 
an acute angle from it. 

On the completion and sealing off of the 
second cell a third lateral shaft with cell is 
excavated almost immediately below the 
first after a slight deepening of the vertical 
shaft. A fourth shaft and cell follows al- 
most immediately below the second in the 



same plane as the third. Excavation of fur- 
ther lateral shafts with cells follows this 
pattern so that a double "stack" of sub- 
horizontal cells (excavated cells each con- 
taining a constructed mud-cell) forms to 
one side of the vertical shaft. 

Of the 22 nests investigated at Mariental 
on 10 March 5 (22.7%) had not yet reached 
the stage of lateral shaft excavation, 4 
(18.2%) had reached lateral shaft excava- 
tion but not cell excavation, 6 (27.3%) had 
one cell each, and 5 (22.7%) had two cells. 
Twelve days later 12 nests were investi- 
gated, 4 (33.3%) had one cell each, 1 (8.3%) 
two cells, 2 (16.7%) three cells, 2 (16.7%.) 
four cells, 1 (8.3%) 11 cells, and 1 (8.3%) 
13 cells. 

Taking two as possibly the largest num- 
ber of cells per nest on 10 March and 13 
twelve days later, it is estimated that it is 
possible that a cell could be prepared, ovi- 
posited into and provisioned in a single 
day. This is of course a very rough esti- 
mate but would be comparable with the 
rate observed for Celonites latitarsis Gess 
(Gess and Gess 1992) and Masarina strucki 
Gess (Gess, Gess and Gess 1997). 

Life histon/. — From two nests each with 
a constructed mud-cell, empty except for 
an egg, it was possible to establish the po- 
sitioning of and appearance of the egg. 
The eggs were white, slightly curved, 
more rounded at one end than at the oth- 
er, 2.6 and 2.22 mm in length and 0.72 and 
0.68 mm in diameter at mid-length. Each 
egg was positioned across the inner end 
of the mud-cell, cemented at its narrower 
end to the mud-cell wall. 

A larva initially feeds only from one 
side of the mass of provision so that the 
"pollen loaf" extracted from a cell with a 
feeding early instar larva is not of even 
diameter along its length, but has an acen- 
tric process at the inner end, the length of 
the process in relation to the length of the 
provision being in proportion to the size 
of the larva. 

The larva consumes the entire provision 
and then defecates at the inner end of the 



Volume 10, Number 1, 2001 



25 



mud-cell. The fecal mass forms a mustard 
coloured deposit on the inner surface of 
the mud-cell to a distance of 3^ mm from 
the inner end. Microscopic examination 
revealed that it contains a mass of empty 
pollen grain walls, in no way macerated, 
with the pores widely extended. In addi- 
tion approximately half-way along the 
length of the mud-cell an irregular mass 
of white crystaline matter is deposited. In 
several of the cells investigated fungal hy- 
phae bearing penicillate sporangiophores 
and white spores covered the fecal mass. 

The mud-cell walls from the edge of the 
fecal layer to approximately 10 mm from 
the inner end of the cell are lined by the 
larva with a parchment-like silken layer. 
At approxiniately 10 mm from the inner 
end of the cell it constructs a seal. Viewed 
from within the mud-cell the seal is in the 
shape of a round, flat-bottomed dish and 
viewed from the mud seal looking into the 
cell it has the appearance of a truncated 
cone. Both surfaces are brown and var- 
nished in appearance. Between the inner 
and outer sloping walls is a series of silken 
parchment-like lamellae (Fig. 23). The cen- 
tral disc which forms the bottom of the 
inner "dish" and outer inverted "truncat- 
ed cone" is thin and translucent. As the 
silken covering of the mud-cell walls only 
extends to the fecal mass there is not a 
complete cocoon. 

On emerging the adult "cuts" out the 
flat, varnished "bottom" of the dish-like 
seal leaving a circular lamellate collar in 
place. The "varnished" disc is left pushed 
to one side. 

VoUinism. — The findings suggest that 
there can be at least two generations of P. 
)iamibiensis per year. Clearly the nests in- 
vestigated on 9 and 10 March were all re- 
cently started by recently emerged fe- 
males. Those investigated on 22 March 
were a mixture of recently started nests 
and nests of which the initiation was prob- 
ably contemporary with those investigat- 
ed on 10 March. This suggested an initial 
staggered emergence of females. The nests 



investigated on 31 March and 2 April, af- 
ter a week of rains, were all recently start- 
ed. These could have been initiated by a 
fresh flush of females of the same gener- 
ation as the nest builders of 10 March or 
a second generation of adults. 

That P. iiamibiensis is possibly bivoltine 
was demonstrated further, when, later in 
April, a male and two females emerged 
from three mud-cells collected on 22 
March from nests of actively nesting fe- 
males, there being no evidence of re-use 
of natal nests. 

DISCUSSION 

Cess and Cess (1980 and 1992) and S.K. 
Gess (1996) discussed possible evolution- 
ary sequences in the Masarinae. They sug- 
gested a sequence from the excavation of 
burrows with excavated cells only (as 
practised by Cermnius damarinus of species 
Group 4, Ceramius species of Group 8 and 
Trimeria howardi) through excavated bur- 
rows with constructed earthen-cells with- 
in excavated cells with earth for construc- 
tion being derived from within the bur- 
row (as practised by Ceramius species of 
groups 2, 3, 5, 6, and 7, at least three spe- 
cies of Paragia and at least two species of 
Jugurtin) to the presumably more ad- 
vanced construction of aerial earthen-cells 
(as typically practised by Celoiiites, Pseu- 
dotriasaris and Gayella). 

That P. iiamibitvisis constructs earthen- 
cells (mud-cells) within excavated cells 
suggests that this behaviour is plesio- 
morphic for Masarinae and that excava- 
tion of cells without constructing cells 
within them is, for Masarinae, derived. 

Descriptions of nesting by Eiiparagia scu- 
tellaris Cresson (Williams 1927, Clement 
and Grissell 1968 and Trostle and Torchio 
1986) do not suggest that Euparagia (Eu- 
paragiinae, formerly included in Masari- 
dae) constructs earthen-cells within exca- 
vated cells. Similarly, there appears to be 
no evidence for construction of cells in 
self-excavated cells by any Eumeninae. It 
would seem that construction of earthen- 



26 



|(H'K\'AI (1F HYMfN'OPIHRA RESEARCH 






23) 

Figs. 19-23. Nest burrow and constructed mud-cells of PriMDimsaris iianiibiciisl^. 19, Burrow cut in vertical 
plane. 20, Constructed mud-cells, four viewed end on (note papillate inner end of mud-cells) and one cut 
transversely to show thickness of ceil wall and smoothed inner surface. 21, Group of constructed mud-cells 
with second cell from top showing "fish-scale" pattern. 22, Group of constructed mud-cells sealed off from 
cut main shaft which is shown cut vertically. 23, Constructed mud-cell cut longitudinally, showing fully grown 
larva and silken parchment-like closure. 



Volume 10, Number 1, 2001 



27 



cells within self-excavated cells is, in the 
Vespidae, probably restricted to Masari- 
nae. Indeed, construction of cells within 
self-excavated cells by aculeate wasps ap- 
pears to be restricted to Masarinae and 
certain Apoidea, the two groups of acu- 
leates that provision with pollen and nec- 
tar. 

The egg of £. scutellaris is, like that of 
Eumenidae, attached to the cell wall by a 
thread (Trostle and Torchio 1986) suggest- 
ing that attachment of the egg by a thread 
is plesiomorphic for Vespidae. It is there- 
fore of note that the egg of P. uamihietisis, 
like that of all other Masarinae, for which 
the egg is known, is not attached by a 
thread. 

S.K. Gess (1996) stated that as a general 
rule at temperate latitudes pollen wasps 
appear to be univoltine but that Zucchi et 
al. (1976) suggested that T. howardi in sub- 
tropical South America may be bivoltine. 
It is therefore of particular interest that it 
has now been demonstrated that P. nami- 
bieiisis is bivoltine. This suggests that the 
recorded second flush of nesting by Jugur- 
tia confusa Richards (Gess and Gess 1980) 
after late summer rain in the Eastern Cape 
of South Africa may indicate that under 
optimal summer conditions Jugurtia may 
also be bivoltine. It is therefore possible 
that whether or not pollen wasps in south- 
ern Africa are uni- or bi-voltine varies 
from year to year with varying spring and 
summer rainfall patterns. 

ACKNOWLEDGMENTS 

Thanks are expressed to the following for much 
appreciated assistance: Estelle Brink of the Schonland 
Herbarium, Albany Museum, Grahamstown, and Co- 
leen Mannheimer and Sabine Austaller, both of the 
National Botanical Research Institute, National Her- 
barium of Namibia, Windhoek, for assistance with 
the identification of the plant specimens; Fred Gess 
of the Albany Museum, Grahamstown, my co-re- 
searcher in the study of aculeate wasps and bees of 
the semi-arid areas of southern Africa, for identifica- 
tion of flower visitors, for taking the photographs re- 
produced as Figs. 3 and 4), and for valuable discus- 
sion and encouragement; Chris Whiteley of the De- 
partment of Biochemistry and Microbiology, Rhodes 



University, Grahamstown for undertaking the sugar 
analysis on the walls of constructeci mud-cells of P. 
iiamibiL'tisis; Erie Grissell and Philip Torchio for their 
constructive reviewing of an earlier version of the 
manuscript; the Namibian Ministry of Environment 
and Tourism for granting a permit to conduct re- 
search and collect biological specimens in Namibia; 
and the South African National Research Foundation 
(formerly Foundation for Research Development) for 
running expenses grants for fieldwork during the 
course of which the present study was undertaken. 

LITERATURE CITED 

Clement, S. L. and E. E. Grissell. 1968. Observations 
of the nesting habits of Eupnrni^in scutcllnris Cres- 
son (Hymenoptera: Masaridae). Pan-Pacific Ento- 
mologist 44: 34-37. 

Gess, F. W. 1998. Priscomasaris nannbicnsis Gess, a new 
genus and species of Masarinae (Hymenoptera: 
Vespidae) from Namibia, southern Africa, with a 
discussion of its position within the subfamily. 
journal of Hymenoptera Research 7: 296-304. 

Gess, F. W. and S. K. Gess. 1990. Ethological studies 
of Jugurtia confusa Richards, Ceramius capicola 
Brauns, C. liiwaris Klug and C. lichtensteinii 
(Klug) (Hymenoptera: Masaridae) in the eastern 
Cape Province of South Africa. Annals of the Cape 
Provincial Museums (Natural History) 13: 63-83. 

Gess, F. W. and S. K. Gess 1992. Ethology of three 
southern African ground nesting Masarinae, two 
Celonites species and a silk spinning Quartinia 
species, with a discussion of nesting by the sub- 
family as a whole (Hymenoptera: Vespidae). 
journal of Hymenoptera Research 1: 145-155. 

Gess, S. K. 1996. The Pollen Wasps: ecology and natural 
history of the Masarinae. Cambridge, Mass.: Har- 
vard University Press. 340 pp. 

Gess, S. K. 1999. Distribution and ethology of Cer- 
amius damarmus Turner (Hymenoptera: Masari- 
nae) in Namibia. In: Byers, G. W., Hagen, R. H. 
and Brooks, R. W. (eds.). Entomological Contri- 
butions in Memory of Byron A. Alexander. Uni- 
versity of Kansas Natural History Museum Special 
Publication 24: 25-32. 

Gess, S. K., F. W. Gess and R. W. Gess. 1997. Update 
on the flower associations of southern African 
Masarinae with notes on the nesting of Masarina 
stucki Gess and Celonites gariepensis Gess (Hy- 
menoptera: Vespidae: Masarinae) in southern Af- 
rica, journal of Hymenoptera Research 6: 75-91. 

Giess, W. 1971. A preliminar)' vegetation map of 
South West Africa. Dinteria 4: 1-114. 

Lovegrove, B. 1993. The Living Deserts of Southern Af- 
rica. Cape Town: Fernwood Press. 224 pp. 

Rutherford, M. C. and R. H. Westfall. 1986. Biomes 
of Southern Africa— an objective categorization. 
Memoirs of the Botanical Survey of South Africa 
54: 1-98. 



28 



Journal o¥ Hymenoptera Research 



Torchio, P. F. 1970. The ethology of the wasp, Pseii- 
doiiiasaris cdivanisii (Cresson), and a description 
of its immature forms (Hymenoptera: Vespoidea, 
Masaridae). Coiitrihutitvis in Science 202: 1-32. 

Trostle, G. E. and P. F. Torchio. 1986. Notes on the 
nesting biology and immature development of 
Euparagia scutellaris Cresson (Hymenoptera: Ma- 
saridae). Journal of the Kansas E}itoiiiologicnl Society 
59: 641-647. 

Williams, F. X. 1927. Euparagia scutellaris Cresson, a 
masarid wasp that stores its cells with the young 
of a curculionid beetle. Pan-Pacific Entomologist 4: 
38-39. 

Zucchi, R., S. Yamane and S. F. Sakagami. 1976. Pre- 
liminary notes on the habits of Triineria howanii, 
a Neotropical communal masarid wasp, with de- 
scription of the mature larva (Hymenoptera: Ves- 
poidea). Insecta nmtsuuiarana. new Ser., 8: 47-57. 

APPENDIX 1 

Collection records of Priscoinasaris lunnibiensis Cess 
listed under the vegetation types of Giess (1971) (giv- 
en by number and name, see also Fig. 1) and the bi- 
omes of Rutherford and Westfall (1986) as adapted 
by Lovegrove (1993) (see Fig. 2). 

5 — Mopane Savanna (falling within the Nama Karoo 



Biome) 
20 13 BB [20.15-20.30S 13.45-14.00E], Wereldsend 

(M. Penrith, v. 1982) (pers. com. ]. Carpenter, 

29.x. 1999) 
20.17S 14.05E, between Palm and Khorixas (F. W. 

and S. K. Gess, 31.iii.l997) 
20.26S 14.54E, 15.5 km by road west of Khorixas (F. 

W. and S. K. Gess, 31.iii.l997) 
20.31S 14.56E, 23 km by road from Khorixas to Uis 

(F. W. and S. K. Gess, l.iv.l997) 

9 — Dwarf Shrub Savanna (falling within the Nama 
Karoo Biome) 
24.1 IS 16.56E, southeast of Bullsport (F. W. and S. 

K. Gess, ll.iii.2000) 
24.40S 17.57E, 5 km south of Mariental by road to 

Keetmanshoop (F. W. and S. K. Gess, 10 and 

31.iii.2000 and 2.iv.2000) 
24.58S 17.55E, 43 km south of Mariental by road to 

Keetmanshoop (F. W. and S. K. Gess, 4.iv.l997) 
25.17S 17.50E, 7 km south of Gibeon railway siding 

by the Mariental/ Keetmanshoop road (F. W. 

and S. K. Gess, 9.iii. and 3.iv.2000) 
25.24S 17.54E, 97 km south of Mariental by road to 

Keetmanshoop (F. W. and S. K. Gess, 4.iv.l997) 
25.53S 18.07E, Tses, 161 km south of Mariental by 

road to Keetmanshoop (F. W. and S. K. Gess, 

4.iv.l997 and 17.iv.l998) 



]. HYM. RES. 
Vol. 10(1), 2001, pp. 29-54 

The Australian Species of Pachyneuron Walker 
(Hymenoptera: Chalcidoidea: Pteromalidae) 

Gary. A. P. Gibson 

Systematic Entomology Section, Eastern Cereal and Oilseed Research Centre, Agriculture and 
Agri-Food Canada, Ottawa, Ontario, Canada KIA 0C6 (E-mail: gibsong@em.agr.ca) 



Abstract. — Four species of Pncln/iiciiwii Walker are recognized from Australia: P. aphidis (Bouche 
1834), P. emersoni Girault (1916), P. nclsoni Girault (1928) and P. rieki Gibson, n. sp. A lectotype 
is designated for P. emersoni. Pachyneuron kingsleyi Girault (1916) is formally synonymized with 
P. emersoni (new synonymy). Males and females of the four species are differentiated by key 
featvires, illustrated, and compared with morphologically similar species present in other regions. 
Pachyneuron oiicrsoni and P. rieki are restricted to Australia, P. aphidis and P. iielsoiii are more 
widely distributed. World distribution is summarized for P. aphidis and P. ne!so)ii and Australian 
distribution and host records are compiled for all the species. 



Pachyneuron Walker consists of about 50 
recognized world species with the follow- 
ing distribution as listed by Noyes (1998): 
Afrotropical (4), Australasian (5), Oriental 
(8), Neotropical (11), Nearctic (12) and Pa- 
learctic (28). Szelenyi (1942) gave a key to 
the Palearctic species, Graham (1969) to 
the European species and Kamijo and 
Takada (1973) to the Japanese species, but 
in other areas the species are unrevised 
and some distributional records listed in 
Noyes (1998) are questionable. Most spe- 
cies are hyperparasites of Aphididae or of 
other plant sucking Hemiptera (Coccoi- 
dea, Psylloidea) through their Braconidae 
(Ichneumonoidea) or Aphelinidae and En- 
cyrtidae (Chalcidoidea) primary parasit- 
oids, or are primary parasitoids or hyper- 
parasitoids of the predators of these plant 
pests (Diptera: Syrphidae, Chamaemyi- 
idae; Coleoptera: Coccinellidae; Neurop- 
tera: Chrysopidae). Some species are also 
recorded as pupal parasitoids of mining or 
gall forming Diptera (Agromyzidae, Chlo- 
ropidae, Cecidomyiidae) or as egg para- 
sitoids of several families of Lepidoptera 
(apparently as hyperparasitoids), and 
there are rare records from other families 



of Diptera, Hymenoptera and Coleoptera 
(Noyes 1998). 

Boucek (1988) listed four species of Pa- 
chyneuron from Australia, but suggested 
that P. kingsleyi Girault was probably only 
a form of P. emersoni Girault and estimat- 
ed that there were probably five valid spe- 
cies. Based on the very few localities listed 
by Boucek (1988) for the species and the 
absence of other than the original publi- 
cations of Girault on Australian Pachyneu- 
ron, the genus might be thought to be rel- 
atively rare and unimportant. However, 
three of the four recognized species are 
common and two are widely distributed 
throughout Australia (Figs. 49-51). I ex- 
amined over 2,000 specimens for this 
study and the species undoubtedly are 
major factors in the population dynamics 
of Australian aphids and their syrphid 
predators. The purpose of this study is to 
differentiate the Australian species and to 
tabulate the known hosts anci distribution 
of the species in Australia. 

MATERIALS AND METHODS 

Literature citations for W.H. Ashmead 
and A. A. Girault incorporate the paper 
numbers, between brackets following the 



30 



Journal of Hymenoptera Research 



year of publication, that are used in their 
bibliographies by Crawford (1908) and 
Dahms (1978), respectively. Morphologi- 
cal terms and abbreviations used for struc- 
tures mostly follow Gibson (1997). Newly 
used abbreviations and terms are: 'mvw' 
for 'marginal vein width', the maximum 
width of the marginal vein, and petiole 
'body' (Fig. 22) for the more or less rect- 
angular portion posterior to the constrict- 
ed or tapered petiole 'neck' (Fig. 22) that 
articulates with the propodeal foramen. 
Measurements were made from dry- 
mounted specimens using an ocular mi- 
crometer with 100 divisions per centime- 
tre and a binocular microscope with zoom 
magnification up to 225 X. Specimens for 
scanning electron microscopy (SEM) were 
prepared following Bolte (1996); illustra- 
tions of P. aphidis were made from speci- 
mens from North America. The SEM mi- 
crograph negatives were converted into a 
digital format using a 35mm scanner. Pho- 
tographs of forewings mounted in Canada 
Balsam on slides were taken using a digital 
camera mounted on a dissecting micro- 
scope. These digital images were enhanced 
using Adobe Photoshop®, and assembled 
into final plates using CorelDraw®. Distri- 
bution maps were generated using Bio- 
link© . Only those localities whose position 
could be determined unequivocally were 
mapped so that the maps generally are 
less comprehensive than the listed rec- 
ords. Length of the sections summarizing 
material examined under 'Distribution' for 
each species was reduced using the fol- 
lowing procedures: all specimens validat- 
ing locality records are in ANIC unless 
otherwise indicated; locality records are 
listed in alphabetical order with different 
localities separated by a period, records 
with the same primary locality are sepa- 
rated by semicolons and the primary lo- 
cality is omitted from the second and sub- 
sequent records; the sex and number of 
specimens examined are not given for the 
three common species; all collection dates 
have been standardized, including omit- 



ting the first two numerals of the year; and 
the four most frequent collectors, C.J. Bur- 
well, J.C. Cardale, I.D. Naumann and J.S. 
Noyes are shortened to CJB, JCC, IDN and 
JSN, respectively. The study was based on 
specimens provided by the individuals 
and collections listed below; acronyms are 
used in the text to denote depositories of 
specimens; those collections denoted with 
an asterisk provided type material or oth- 
er specimens of Nearctic and Palearctic 
species that were used to help establish 
correct nomenclature. 
ANIC Australian National Insect Col- 
lection, CSIRO, Canberra, ACT 
(J. Cardale and S. Schniidt) 
ASCU Agricultural Scientific Collec- 
tions Unit, Orange Agricultural 
Institute, Orange, NSW (M. 
Fletcher and P. Gillespie) 
BMNH'^ The Natural History Museum, 

London, England (J. Noyes) 
CNCP Canadian National Collection 
of Insects, Ottawa, ON, Canada 
DPIQ Queensland Department of Pri- 
mary Industries, Brisbane, QLD 
(J. Donaldson) 
HFES* Hokkaido Forest Experiment 
Station, Bibai, Hokkaido, Japan 
(K. Kamijo) 
MHNG* Museum d'Histoire naturelle, 
Geneva, Switzerland (B. Merz) 
QMBA Queensland Museum, Brisbane, 

QLD (C. Burwell) 
UQIC University of Queensland Insect 
Collection, St. Lucia, QLD (G. 
Daniels) 
USNM* United States National Ento- 
mological Collection, U.S. Na- 
tional Museum of Natural His- 
tory, Washington, DC (E. Gris- 
sell) 
WARI Waite Agricultural Research In- 
stitute, University of Adelaide, 
Glen Osmond, SA (A. Austin) 

Pachxjneiiron Walker 

Pachy neuron Walker, 1833: 371, 380. Type spe- 
cies: Pachifiicuron fonnosuni Walker, by nion- 
otypy. 



Volume 10, Number 1, 2001 



31 



Pachynevron Blanchard, 1840: 260, 266. Unjusti- 
fied emendation. 

Pachynevrum Agassiz, 1846: 778. Unjustified 
emendation. 

Serimus Brethes, 1913: 90. Type species: Serimus 
argcntinus Brethes, by monotypy. Synonymy 
by De Santis, 1957: 118. 

Propachyneuronia Girault, 1917[327]: 102. Type 
species: Encyrtus siphoiophorae Ashmead, by 
original designation. Synonymy by Gahan, 
1918: 66. 

Nepnchyucuron Girault, 1917[330]: 9. Type spe- 
cies: Pachyneuron eras Girault, by original 
designation. Synonymy by Timberlake, 1926: 
308. 

Eupachifiieuwii Blanchard in Leiboff, 1948: 256. 
Type species: Eupachyneuron bosqui Blan- 
chard, by monotypy. Synonymy by Boucek, 
1988: 441. 

Atrkhoptilus Delucchi, 1956: 141-142. Type spe- 
cies: Pachyneuron aeneum Masi, by original 
designation. Synonymy by Boucek, 1965: 18. 

Pachyneuron {Serimus); De Santis, 1975: 8-10. 
Change of status. 

Diagnosis. — Head without distinct malar 
depression; clypeus with apical margin 
shallowly emarginate (Figs. 17, 18) to pro- 
duced (Figs. \, 2); gena and lower face ex- 
cluding clypeus mostly isodiametric-retic- 
ulate (Fig. 18); torulus at or above lower 
orbit near middle of face (Figs. 1, 13, 16, 
17, 25, 30, 47). Mandible with four teeth 
(Figs. 1, 2). Antenna 13-segmented with 2 
or 3 anelli; scape of female, when ap- 
pressed to head, extending to anterior 
ocellus; scape of male subequal in width 
or evenly tapered to apex (Figs. 9, 21, 33, 
34, 45, 46). Pronotum visible in dorsal 
view, with pronotal carina (in regional 
species) (Figs. 3, 4, 14, 26, 37, 38). Meso- 
notum reticulate, the sculpture formed by 
raised ridges; mesoscutum with incom- 
plete notauli (Figs. 3, 4); mesopleuron 
with upper mesepimeron shiny and much 
more finely sculptured than lower mese- 
pimeron (Figs. 4, 38). Propodeum with su- 
pracoxal flange shorter than length of nu- 
cha (Fig. 15). Metacoxa bare dorsobasally 
(Figs. 4, 38), outer surface smooth to cori- 
aceous-reticulate, much more finely sculp- 



tured than reticulate femoral depression 
(Figs. 4, 38). Forewing with marginal vein 
noticeably thicker than stigmal or post- 
marginal veins and at least slightly wid- 
ened distally, about as long as stigmal 
vein and at most 0.35 length of costal cell 
(Figs. 6, 24, 36, 48). Caster variably dis- 
tinctly petiolate (Figs. 11, 22, 27, 41); first 
gastral sternum with anterior margin un- 
modified, not produced into flange be- 
neath petiole (Figs. 12, 28, 42); terga flat to 
low convex in critical-point dried female, 
often flat or collapsed in air-dried female. 

Remarks. — Australian Pachyneuron can 
be identified to genus using the key of 
Boucek (1988). Individuals are most likely 
to be confused with specimens of the 
monotypic genus Inkaka Girault (Boucek 
1988, figs. 767-769), but specimens of /. 
quadridentata (Girault) differ conspicuous- 
ly by lacking a carinately margined pron- 
otal collar, the pronotum being almost 
vertical and not visible in dorsal view. In- 
dividuals of Inkaka also have an obvious 
malar depression, the antennal toruli 
slightly below the level of the lower orbits, 
and a more elongate-slender marginal 
vein that is at least 0.4 times as long as the 
costal cell; in females the scape does not 
extend to the anterior ocellus, and in 
males the scape has two distinct subapical 
lobes on its anterior outer margin so as to 
appear emarginate subapically. 

Coruna Walker and Euneura Walker are 
not yet recorded from Australia, but com- 
prise species that are hyperparasites of 
aphids and that are morphologically sim- 
ilar to species of Pacinjneuron. It probably 
is only a matter of time before species of 
one or both genera are accidentally intro- 
duced into Australia. Individuals of Co- 
runa have sulcate notauli that extend to 
the transscutal articulation and therefore 
are easily distinguished from Pachyneuron; 
more subtle features differentiate Euneura 
from Pachyneuron. In Euneura the lower 
face is more extensively longitudinally 
striate-reticulate (Boucek 1988) and the su- 
pracoxal flange is longer than in Pachyneu- 



32 Journal of Hymenoptera Research 

ron (Kamijo and Takada 1973). Also, in Eu- Females of Eimeura also have the metaso- 
neum the metacoxa and femoral depression ma strongly convex and hence more sub- 
are similarly reticulate, whereas in Pachy- circular in cross section than do females of 
neuron the metacoxa is much more finely Pachyneuron, but this difference is less ob- 
sculptured than is the femoral depression, vious in critical-point dried individuals. 



KEY TO AUSTRALIAN SPECIES OF PACHYNEURON WALKER 

1 Both sexes: propodeum uniformly coriaceous anterior to nucha, without plical furrow or 
ridges (Fig. 5); petiole body in dorsal view shiny, virtually smooth and strongly trans- 
verse (Fig. 11); clypeus medially convex and apically rounded to angulate (Figs. 1, 2). 
Female: flagellum with 3 anelli and 5 funicular segments (Figs. 7, 8). Male: anteiina brown 

except possibly for extreme base of scape and legs with femora mostly infuscate 

P. aphidis (Bouche) 

- Both sexes: propodeum with variably distinct, more or less W-shaped complex of plicae 
and costulae anterior to nucha (Figs. 15, 29, 39); petiole body in dorsal view strongly 
reticulate to reticulate-rugose and often longer than wide (Figs. 22, 27, 41); clypeus 
medially flat to depressed and shallowly emarginate (Figs. 17, 18). Female: flagellum 
with 2 anelli and 6 funicular segments (Figs. 19, 20, 31, 32, 43, 44). Male: antenna with 

at least scape mostly or entirely yellow and legs yellow 2 

2(1) Female: flagellum clavate with longitudinal sensilla along almost entire length of flagellar 

segments and with adpressed setae (Figs. 19, 31, 43) 3 

- Male: flagellum filiform with longitvidinal sensilla in apical half of flagellar segments 
and with semierect setae (Figs. 21, 33, 45) 5 

3(2) Forewing with basal cell separated from speculum by oblique line of at least 7 setae on 
dorsal surface of basal fold and with 2 or more setae within basal cell near apex (Fig. 
48); petiole body with 1-3 setae projecting anterolaterally from each side near middle 
(Fig. 41) P. rieki Gibson, new species 

- Forewing usually completely bare basally, but at most with 1 or 2 setae on dorsal surface 
of basal fold (Figs. 23, 24, 35, 36); petiole body without setae projecting from lateral 
margin (Figs. 22, 27) 4 

4(3) Forewing without marginal fringe (Figs. 35, 36); marginal vein comparatively short and 
thick, length less than 2.5 times maximum width (Fig. 36), and postmarginal vein only 
slightly (less than 1.25 times) longer than stigmal vein (Fig. 36); propodeum reticulate- 
coriaceous anterior to nucha, similarly or even more strongly sculptured medially than 
laterally (Fig. 29) P. nelsoni Girault 

- Forewing with marginal fringe (Figs. 23, 24) and /or with relatively elongate-slender 
marginal vein at least 3.5 times as long as wide, and with postmarginal vein distinctly 
(at least 1.5 times) longer than stigmal vein (Fig. 24); propodeum with comparatively 
shiny, finely coriaceous to virtually smooth pentagonal or hexagonal region anterior to 
nucha (Fig. 15) P. emersoni Girault 

5(2) Forewing without marginal fringe (Figs. 35, 36); flagellar segments oblong, the middle 
segments less than 1.8 times as long as wide (Fig. 33); antenna uniformly yellowish or 

with flagellum light brown P. nelsoni Girault 

Forewing with marginal fringe (Figs. 23, 24, 48); flagellar segments elongate, middle 
segments more than 1.8 times as long as wide (Figs. 21, 45); antenna usually with dark 
brown flagellum contrasting distinctly with yellow scape 6 

6(5) Forewing with basal cell separated from speculum by oblique line of at least 7 setae on 
dorsal surface of basal fold (Fig. 48); head with lower face uniformly convex (Fig. 47); 

scape in profile with line of distinct setae along anterior margin (Fig. 46) 

P. rieki Gibson, new species 



Volume 10, Number 1, 2001 



33 



- Forewing with basal cell and speculum uniformly bare (Figs. 23, 24) and /or head with 
lower face distinctly depressed or concave lateral to convex supraclypeal area and clyp- 
eus (Fig. 16); scape in profile without line of distinct setae along anterior margin (Fig. 
21b) P. eniersoni Girault 



Pachyneuron aphidis (Bouche) 

(Figs. 1-12, 49) 

Dip^lolcpis Aphidis Bouche, 1834: 170. Syntypes; 
types lost according to Graham, 1969: 842. 
Sex described: both. 

Pteromalus miniitissimiis Forster, 1841: 28. Lec- 
totype male, designated by Delucchi, 1955: 
138. Type data: Germany, bred on Aphidiiis 
rosarum. Type depository: Naturhistorisches 
Museum, Vienna. Sex described: male. Syn- 
onymy by Reinhard, 1859: 195. 

Pachyneuron pruni Walker, 1850: 128. Lectotype 
female, designated by Graham, 1969: 842. 
Type data: Prussia. Type depository: BMNH. 
Sex described: female. Synonymy by Gra- 
ham, 1969: 842. 

Pachyneuron aphidis; Reinhard, 1859: 195. 
Change of combination. 

Encyrtus siphonophorae Ashmead, 1886[36]: 131. 
Syntypes, female (examined). Type data: 
USA, Florida [Jacksonville] bred in 1881 from 
orange aphis {Siphonophora citrifolii Ash- 
mead). Type depository: USNM, type no. 
4860. Sex described: female. Synonymy by 
Boucek, 1988: 441, 442. 

Pachyneuron aphulivora Ashmead, 1887[37]: 14. 
Syntypes, both sexes (examined). Type data: 
USA, Florida [Jacksonville], bred June 6 from 
the cabbage aphis (Ap>}ns brassicae L). Type 
depository: USNM, type no. 2854. Sex de- 
scribed: female. Synonymy with E. siphono- 
phorae by Girault, 1917[327]: 102. Note: Ac- 
cording to Timberlake (1918: 402) Girault's 
synonymy is incorrect because notes of A.B. 
Gahan on the types, "taken when they were 
in a better state of preservation than at pre- 
sent, show that aphidivorum has only two 
ring-joints". However, although Ashmead 
described only females, both females and 
males are labelled as syntypes in the USNM 
collection and Gahan's note undoubtedly re- 
ferred to a male. 

Pachyneuron maidaphis Ashmead, 1888: 23. Syn- 
types, female (examined). Type data: USA, 
Florida [Lake City]. Type depository: USNM, 
type no. 26530. Sex described: female. Syn- 



onymy with £. siphonophorae by Girault, 
1917[327]: 102. 

Pachyneuron micans Howard, 1890: 246. Syn- 
types, female (examined). Type data: USA, 
Indiana, Lafayette, reared by Webster from 
Siphonophora avenae. Type depository: 
USNM, type no. 1467. Sex described: both. 
Synonymy with E, siphonophorae by Girault, 
1917[327]: 102. 

Pachyneuron gifuensis Ashmead, 1904[243]: 158. 
Syntypes, female (examined). Type data: [Ja- 
pan], Gifi.1, bred by Y. Nawa [Oct. 1902] from 
an Aphis. Type depository: USNM, type no. 
7190. Sex described: female. Synonymy by 
Kamijo & Takada, 1973: 57. 

Serinius argentinus Brethes, 1913: 91. Holotype 
male. Type data: Argentina: Buenos Aires. 
Sex described: male. Synonymy by De Santis, 
1957: 119. 

Pachyneuron lali Mani, 1939: 81. Holotype fe- 
male, by original designation. Type data: In- 
dia: Karnal (Punjab), 15.iii.l938, K. B. Lai, 
bred from Aphis runiicis L. on Solanum ni- 
grum. Type depository: Indian Agricultural 
Research Institute, New Delhi. Sex described: 
female. Synonymy by Boucek et ai, 1978: 451. 
Note: purported description of male applies 
to female based on description of three anelli, 
see Boucek et al. (1978). 

Pachyneuron ferrierei Mani, 1939: 83. Syntypes, 
male. Type data: India: Chaubatia, U.P., 
R.N. I., 22.viii.1935, parasitic on an aphid 
causing leaf curl. Type depository: Indian 
Agricultural Research Institute, New Delhi. 
Sex described: male. Synonymy by Boucek et 
ai, 1978: 45. Note: type .series mistakenly 
stated as female (Boucek et al. 1978). 

Eupachyneuron bosqui Blanchard /;/ Leiboff, 
1948: 256. Type status unknown. Type data: 
Argentina, La Pampa, reared from Schyzaphis 
granunum. Sex described: female. Synonymy 
with P. aphidis by Boucek, 1988: 442. 

Pachyneuron minutissimum; Delucchi, 1956: 129, 
137-139. Change of combination. 

Pacliyneuron triarticulata Mani & Saraswat, 1974: 
98-100. Holotype female, by original desig- 



34 



Journal of Hymenoptera Research 



nation. Type dc\Lr. India: Northwest Hima- 
laya, Dalhousie (Ahla catchment area, Kha- 
jjiar), M.S. Mam and party, 18 & 27.V.1971. 
Type depository: USNM, database no. 
0700023. Sex described: female. Synonymy 
by Boucek et al, 1978: 451. 

Pachyneuwn {Serimus) siyihonophorac; De Santis, 
1975: 9. Change of status. 

Pachyiicuroii aphidis; Boucek, 1988: 442. 

Femnle. — Body brown to dark brown 
with variably distinct metallic green lus- 
ter; antenna brown except extreme base of 
scape often yellowish; tegula yellow to 
brown; legs with femora variably darkly 
infuscate except apically yellowish, tibiae 
and tarsi yellowish or with tibiae lightly 
infuscate subbasally. Head with clypeus 
convex and apically rounded to angulate 
(Figs. 1, 2). Flagellum with 3 anelli (Fig. 8) 
and 5 funicular segments (Fig. 7); funicu- 
lar segments subquadrate to oblong and 
with long, conspicuous, decumbent setae 
(Figs. 7, 8); longitudinal sensilla extending 
almost entire length of funicular segments, 
separated from each other by distance 
equal to 2-3 sensillar diameters (Fig. 8). 
Forewing (Fig. 6) with marginal fringe; 
with distinct discal setae; dorsally with 
basal cell apically delineated by oblique 
line of setae directed posterobasally from 
base of parastigma; ventrally with poste- 
rior margin of basal cell often delineated 
by longitudinal cubital setal line, and of- 
ten with one to several setae on ventral 
surface near submarginal vein; speculum 
on dorsal surface open posteriorly; costal 
cell with distinct setae on ventral surface; 
veins with following ratios {ii = 10): smv/ 
mv = 2.70-3.28, mv/mvw = 2.75-3.58, 
pmv/mv = 1.57-2.12, pmv/st = 1.64- 
2.21. Mesonotum with highly convex, rel- 
atively slender scutellum (Figs. 3, 4). Pro- 
podeum (Figs. 3-5) strongly transverse, 
uniformly striate-coriaceous without me- 
dian carina, costula, or plical carina except 
near nucha, but with paramedial trans- 
verse depressions basally; spiracle circular 
to slightly oval. Petiole without setae pro- 
jecting from sides; in dorsal view body 



strongly transverse, shiny and virtually 
smooth (Fig. 11); in ventral view divided 
mediolongitudinally by white membra- 
nous region (Fig. 12). 

Male. — Similar to female except flagel- 
lum (Figs. 9, 10) with 2 anelli and 6 funic- 
ular segments; all segments, except possi- 
bly preclaval segment, longer than wide 
(at most about 1.8 times). 

Distribution.— PncJn/iieuro}} aphidis is a 
cosmopolitan species that Noyes (1988) re- 
corded from over 40 countries and all con- 
tinents except Antarctica. 

Australian distribution (Fig. 49) and 
host records based on label data of ex- 
amined specimens are: Australian Capital Ter- 
ritory: Canberra, 2.xii.46, 18.vi.54, E.F. Riek; coll. 
17.V.89, em. 25.V.89, Diacrcticlla rnpnc ex. Brcvicorync 
brnsfiicac on SisyiiibriiDii officinale, V.F. Eastop; coll. 
10.iii.89, em. 18.iii.89, Aphidiiis soiichi Marshall ex. Hif- 
pc)V)in/ziis lactiicae (L.) on Soiichiis olcmceiis, V.F. Eas- 
top. New South Wales: Glossodia, 20.vi.80, parasite 
of Aphis ncrii, D. James. Katoomba, 2.xii.75, IDN 
(UQIC). Merungle Hill, 2.ii.66, on mature oranges, 
M.I. Nikitin (ASCU). Mootwingee Nat. Pk., Old Moo- 
twingee Gorge, 5-8.xi.84, G.R. Brown & H.M Holmes 
(ASCU). Newport, 18.iii.36, ex. Apliis iwrii (ASCU). 
Rob Roy, 29.V.74, ex. ear of wheat, M. Greening 
(ASCU). Rydalmere, 7.vi.61, L.T. Woolcock; 22.ii.68, 
parasitized Brcvicorync brassicac, J.T. Hamilton 
(ASCU). Sydney, 20. v. 49, ex. pumpkin aphids 
(ASCU). Tamworth, l.xi.79, ex. Trioxys complanatus 
on spotted alfalfa aphid, N. Forrester. Ulladulla, 
18.i.72, ex. aphid on Cakilc cdentiila, M. Gray; 10.ii.81, 
H. lactiicac; 17.ii.81. Warrumbungles Nat. Pk. via 
Coonabarabran, 17.xii.74, IDN (UQIC). Wellington, 
23.ii.54, ex. B. brnssicac. Queensland: Beerwah, 3.5 km 
NW (26.50S 152.56E), CJB (UQIC). Brisbane, Alan 
Fletcher lab., ix.82, ex. galls on Rhopnloun/ia califoniica 
Felt, P. McFadyen (QDPl). Bunya Mts. Nat. Pk. nr. 
Westcott Plain (26.52S 151.34E), 6-7.X.84, IDN & JCC. 
Camp Mt., 1.X.61, ex. Aphis ncrii on Asclcpins CKras- 
stuica, E.N. Marks (UQIC). Cecil Plains, D-Vac SIR- 
ATAC trial in cotton, 1981-82, P.D. Rossiter (QDPI). 
Gatton, 16.V.78, ex. Mcropln/as diviilsana, B. Franz- 
mann (QDPI); 5.V.81 (QDPI); D.P.I. Research Station, 
l-7.ix.81, 7-14.ix.81, 2I-28.ix.81 (QDPI). Helidon, 
30.viii.79, ex. Aphidiiis colcmani on Aphis ncrii, B.A. 
Franzmann (QDPI). Lake Broadwater, 25 km SE Dal- 
by (27.22S 151.06E), 2-3.iv.96, CJB (QMBA). Miles, 1 
km W, Dogwood Ck., 7.X.74, I.D. Galloway (QMBA). 
Mt. Glorious (27.20S 152.46E), 27.ix.95, S.G. Evans 
(QMBA). Russell Pk. nr. Mt. Mowbullan (26.53S 
151.37E), 7.X.84, IDN & JCC. Sanford, x.61, E. War- 
wick. Tenthiil, 15.viii.79, ex. Apliidiiis cok'iiuiiii in 



Volume 10, Number 1, 2001 



35 




Figs. 1-6. Pachyneurou aphidis: 1, head, frontal (9); 2, clypeus and mandibles (9); 3, mesosoma, dorsal ($); 
4, mesosoma, lateral (9); 5, scutellum-propodeum (c5); 6, forewing (9). Scale bars = |xm. Abbreviations; bsl 
= basal setal line, csl = costal setal line. 



36 



Journal of Hymenoptera Research 




( 1 ): 7ci, t'litire, 7b, anelli and fu 



Figs. 7-12. Pncln/iiciiroii nplmlls: 7, antenna 

segments, fl-fl, (?); 9, antenna (j): 9a, entii 

ments, tl ,-fl, (d); 11, petiole, dorsal (V); 12, petiole, ventral (?). Scale bars = |jLm. 



' ' -' ■ -r "■- -"-. ..-.nicular segments; 8, basal flagellar 

e, yb, middle funicular segments, fl,-fl,; 10, basal flagellar seg- 



VoLUMH 10, Number 1, 2001 



37 



Apliis iicrii, B.A. Franzmann (QDPI). Toowoomba, 
16.V.75, ex. mummies on Rhopalosiphuin iiiaidis, H. Bri- 
er (QDPI); 15.x. 79, ex. Apliidiits Lolcinnui on Apliis iicrii, 
B.A. Franzmann (QDPI). South Australia; Adelaide, 
Waite Institute, iv.88, hyperparasite of Apliidius soii- 
chi, D. Martin (WARI). "Brecon", 10 km S Keith, 
26.i.82, A.D. Austin (QDPI). Glen Osmond, Waite Ag. 
Res. Inst., coll. 24.iii.82, em. 30.iii.82, ex. pea aphid, 
D. Samoedl; W.A.R.I., 21.i.81, ex. Tnoxxp, SAA culture 
on lucerne, D. Samoedl; Waite Institute, vi.73, ex. 
Brcvicoryiw brassicae and Mi/zus persicae, C. Crawford. 
Pinnaroo, 25 km SSW (35.28S 140.47E), 20.X.89, 
24.X.89, IDN & JCC; 49 km SW (35.42S 140.49E), 
20.X.83, 24.X.83, IDN & JCC. Victoria: nr. Benalla, 
9.ii.78, ex. T. ninculata. 

Hosts.— Noyes (1988) listed 115 species 
and 62 genera as hosts for P. aphidis in the 
following taxa: Diptera (Cecidomyiidae, 
Syrphidae), Hemiptera (Aphidoidea: 
Aphididae, Pemphigidae; Coccoidea: Coc- 
cidae, Kermesidae, Pseudococcidae; Psyl- 
loidea: Psyllidae), and Hymenoptera 
(Chalcidoidea: Aphelinidae, Encyrtidae; 
Ichneumonoidea: Braconidae). Based on 
label data, Australian primary and sec- 
ondary hosts include Aphididae: Acyrtho- 
siphon pisum (Harris), Aphis nerii (Fonsco- 
lombe), Brevicoryne brassicae (L.), Nasonovia 
(= Hyperomyzus) lactucae (L.), Myzus per- 
sicae (Sulzer), Therioaphis maculatn (Buck- 
ton) and Braconidae: Aphidius colemani Vi- 
ereck, Aphidius sonchi Marshall, Diaeretiella 
rapae (Mcintosh), Trioxys complanatus 
(Perez). There is also a single record from 
Merophyas divulsana (Walker) (Lepidop- 
tera: Tortricidae) and an anomalous re- 
cord of 'galls' on Rhopalomyia californica 
Felt. 

Remarks. — Pachyneiiron aphidis is the 
only species of Pachyneiiron in Australia 
with a convex, apically rounded or angu- 
late clypeus (Fig. 2). It is further differen- 
tiated from P. nelsoni and P. emersoni fe- 
males by the presence of a basal setal line 
on the forewing (Fig. 6); however, speci- 
mens of P. rieki (Fig. 48) and rare P. emer- 
soni males also have a forewing basal setal 
line. Females of P. aphidis are also unique 
within the genus because they have 3 
anelli and 5 funicular segments, and the 
flagellum differs from those of other Aus- 



tralian species because it has conspicuous 
decumbent setae similar to males (Figs. 7, 
8). Males are easily distinguished by their 
brown antennae, males of the other spe- 
cies have at least the scape yellow. 

Because of a similar clypeus and pro- 
podeum, individuals of P. aphidis are mor- 
phologically most similar to P. californicum 
Girault (1917[322]), known only from 
America north of Mexico. However, in P. 
californicum the petiole is completely scler- 
otized ventrally (i.e., forming a complete 
tube, cf. Fig. 28) even though short as in 
P. aphidis, and the speculum is usually 
closed on the ventral surface by a line of 
setae along the cubital fold. 

Pachyneiiron emersoni Girault 

(Figs. 13-24, 50) 

Pachyneiiron emersoni Girault, 1916[274]: 229- 
230. Lectotype female, complete (examined), 
here designated: "878 ", "Swan Riv, W. 
Austr.", "G. Compere Collector", " 9 Lecto- 
type Boucek 1985". Type depository: USNM, 
type no. 19691. Paralectotypes, here desig- 
nated: 1 point with mesosoma, same data as 
lectotype (USNM type no. 19691); 1 slide 
with parts of two male antennae under one 
cover slip and a crushed head, one female 
hind leg and two male hind legs under an- 
other cover slip (USNM type no. 19691); 1 
point with mesosoma, same data as lectotype 
(QMBA type no. HY 3568); 1 point with pair 
of middle legs, same data as lectotype plus 
"from Icerya, California, Alex. Craw, import 
from Australia, G. Compere, July 1900" 
(QMBA type no. HY 3568). Notes: Although 
Boucek labelled the two USNM point- 
mounted specimens as lectotype and para- 
lectotype he did not validate these designa- 
tions through publication. Girault's original 
description stated that the USNM has "two 
females on tags plus the slide"; however the 
point-mounted USNM paralectotype is a 
male, based on leg structure and color. The 
male antennae on the slide may belong to 
this specimen but it is unknown to which 
specimen the other parts belong. 

Pachyneiiron kingslcyi Girault, 1916[274]: 230. 
Holotype female (examined). Type data: 
Australia: N.S.W., Brooklyn, 31 October 1914. 



38 



Journal of Hymenoptera Research 



Type depository: QMBA, type no. HY 3569. 

Sex described: female. New synonymy. 
Pachyneuron emersoni; Dahms, 1983: 246; Bou- 

cek, 1988: 442. 
Pachyneuron kingslei/i; Girault, 1927[416]: 335; 

Girault, 1929[431]: 319; Dahms, 1984: 738- 

739; Boucek, 1988: 442. 

Female. — Body dark with variably dis- 
tinct metallic green luster; antenna dark 
brown except with basal half to all of 
scape yellow; tegula yellow; legs with 
femora variably darkly infuscate except 
apically yellowish, tibiae and tarsi yellow- 
ish. Head with clypeus flat to slightly de- 
pressed and apically shallowly emarginate 
(Fig. 13). Flagellum compact-clavate, with 
2 anelli (Fig. 20) and 6 funicular segments 
(Fig. 19); funicular segments slightly lon- 
ger than wide basally to quadrate or 
slightly transverse apically and with ad- 
pressed setae (Figs. 19, 20); longitudinal 
sensilla extending almost entire length of 
funicular segments, separated from each 
other by distance equal to 1-2 sensillar di- 
ameters (Fig. 20). Forewing (Figs. 23, 24) 
usually with marginal fringe; with distinct 
discal setae; dorsally without line of setae 
differentiating apex of basal cell from 
speculum (very rarely with 1 or 2 setae on 
basal fold); ventrally without line of setae 
along cubital fold; costal cell with distinct 
setae on ventral surface (Fig. 23b); veins 
with following ratios {n = 10): smv/mv = 
2.94-3.60, mv/mvw = 3.63^.86, pmv/mv 
= 1.32-1.64, pmv/stv = 1.55-1.80. Meso- 
notum with relatively low convex, broad 
scutellum (Fig. 14). Propodeum (Fig. 15) 
with posteriorly convergent, carinately 
margined plical ridges and usually less 
distinct, often irregularly fl-shaped anter- 
omedian carina or ridge (costula) near 
base, the ridges together differentiating a 
more or less W-shaped basal region with 
coriaceously sculptured anterolateral de- 
pressions from a mostly shiny and smooth 
to finely coriaceous pentagonal or hexag- 
onal posteromedian region anterior to a 
coriaceous or medially smooth and shiny 
nucha, with the short region anterior to H- 



shaped ridge crenulate and the surface lat- 
eral to plical ridges finely coriaceous; spi- 
racle distinctly oval. Petiole without setae 
projecting from sides (Fig. 22); in dorsal 
view distinctly (at least 1.75 times) longer 
than wide, with body slightly to distinctly 
longer than wide and uniformly reticulate 
(Fig. 22); in ventral view completely scler- 
otized with median furrow, the body dis- 
tinctly longer than wide, finely coriaceous- 
reticulate and shiny. 

Male. — Similar to female except as fol- 
lows: body brighter metallic green or blu- 
ish green; legs uniformly bright yellow be- 
yond coxae except metafemur sometimes 
lightly infuscate; head with lower face dis- 
tinctly concave (Fig. 16) to very shallowly 
depressed (Figs. 17, 18) lateral to convex 
supraclypeal area and clypeus; scape en- 
tirely yellow or brownish apically, slightly 
expanded and flattened basally immedi- 
ately above radicle, tapered apically and 
slightly curved, and in lateral view with- 
out distinct line of setae along anterior 
margin (Fig. 21b); flagellum usually dark 
brown, filiform; funicular segments elon- 
gate, middle segments more than 1.8 times 
as long as wide and all segments with 
very sparse longitudinal sensilla within 
apical half and with conspicuous, semi- 
erect setae about as long as width of seg- 
ment (Figs. 21a, c); forewing always with 
marginal fringe; basal cell on dorsal sur- 
face sometimes delineated apically by line 
of up to 7 setae, with 1-3 setae sometimes 
also delineating extreme posteroapical an- 
gle of cell, and rarely with 1-3 setae with- 
in cell toward apex. Petiole usually with- 
out setae, rarely with single seta projecting 
anterolaterally from one side near middle. 

Distrilmtion (Fig. 50). — Australian Capital 
Territory: Bendora, 14.xii.60, D.H. Colless. Blundell's, 
26.ix.30, 14.iv.31, L.F. Graham; 6.i.61, E.F. Riek. Brin- 
dabella Ra., Lees Spring (35.22S 148.49E), 24.xi.81, 
IDN; nr. Lees Spring, 24.xi.31, L.F. Graham; Mt. Gin- 
ini, 24.xi.81, IDN. Canberra, 14.ii.39, from cabbage, 
T.G. Campbell; 3.xii.39; 29.ix.46, 2.X.46, 6.viii.47, 
15.vii.48, 8.xi.48, E.F. Riek; 25.xi.65, O.W. Richards; 
10.V.66, ex. Aphis crnccivora, D. Morgan; 18.i.80, on Eu- 
calyptus blossom, IDN; 20.i.80, on Bncckia blossom, 



Volume 10, Number 1, 2001 




Figs. 13-18. Pachi/nciiwu cmcrsom: 13, head, frontal (9); 14, mesosoma, dorsal ($); 15, scutellum-propodeum 
(9); 16, head, frontal (c5); 17, head, frontal (S); 18, lower face (S) (arrows point to regions of finer .sculpture 
within depressions). Scale bars = [xm. Abbreviations: cos = costula, nuc = nucha, pic = plica, scf = supracoxal 
flange. 



40 



Journal of Hymenoptera Research 




Figs. 19-24. Piicln/iiciimii niursoni: 19, antenna (9): 19a, entire, 19b, anelli and funicular segments; 20, basal 
flagellar segments, f\-i\, (','); 21, antenna (c^): 21a, entire, 21b, scape, 21c, middle funicular segments, fl,-fl^; 
22, petiole, dorsal (V); 23, forewing, SEM of dorsal surface: 23a, entire, 23b, submarginal vein and costal cell 
(arrow points to costal setae) (6); 24, forewing, photograph (9). Scale bars = (xm. 



Volume 10, Number 1, 2001 



41 



IDN. Canberra, nr. airport, 7.iv.53, 14.iv.53, ex. syr- 
phid pupa Eucalyptus iiicIUodora, E. Lewis; 8.iv.53, ex. 
syrphid pupa under Euc. umculosn, E. Lewis. Canber- 
ra, Black Mt., 22.V.53, ex. syrphid pupa Euc. hlakchji, 
E. Lewis; 14.iii.67, 9-16.xi.79, 17-26.xi.79, D.H. Col- 
less; 2-10.iv.68, 28-29.iv.68, light trap; 18.xi.79, Z. Lie- 
pa; 600m, 7-12.iii.80, dry sclerophyll, A. Newton & 
M. Thayer; iii, iv, v, vii, ix, x, 24.x-l.xi, xi.82, IDN & 
JCC. Canberra, Capital Hill, coll. 21.\.53, em. 21.xi.53, 
ex. syrphid pupa Euc. blakclyi, F. Wheelhouse; ex. 
curled leaf £. luclliodorn, 9.iii.53, E. Lewis. Canberra, 
Farrer, 27.xi.80, 7.xii.80, R. Rentz. Gibraltar Falls, 
27.i.84, IDN. Gingera, 6.iii.52, E.F. Riek. Ginninderra, 
22.iii.66, G. Grant. Molonglo, 8.iv.83, ex. syrphid 
pupa under Euc. maculosa, E. Lewis. Mt. Coree, 
30.i.64, D.H. Colless; 2 mi E, 19.xi.68, ]CC & S. Curtis. 
Mt. Franklin, 4 km N (32.27S 148.46E), 21.ii-10.v.77, 
D.C.F. Rentz. Mt. Gingera, 4.ii.65, D.H. Colless; 
13.i.69, Z. Liepa; 21.ii.79, E.C. Zimmerman; 4.i.79, ex. 
lichen, R. Bartell. Picadilly Circus (35.22S 148.48E), 
1240m, xii.84, J. Lawrence, T. Weir & M.L. Johnson; 
(35.22S 148.49E) 24.xi.81, JCC; 17 km SW (35.27S 
148.48E), 24.xi.81, IDN & JCC; 3 km E, Blundell's Ck 
(35.22S 148.50), 850m, ii.84, xii.84. Weir, Lawrence & 
Johnson; ii.87, D.H. Colless; 6 km NW, Wombat Ck 
(35.19S 148.51E), 750m, xii.84, i.85. Weir, Lawrence & 
Johnson. Quenbeyan, 9.iii.53, ex. curled leaf E. uwl- 
liodera, E.L. Raymond; 2.7 km NE, 670m, 5.iv.80, I.F.B. 
Common. Smokers Flat, 6 km E Corin Dam, 4.iv.80, 
J.F. Lawrence. Woods Reserve nr. Gibraltar Falls, 
27.i.84, IDN. New South Wales: Alpine Creek, Kian- 
dra, 9.xii.64, E.F. Riek. Arcadia, 10.ii.67, M.I. Nikitin 
(ASCU). Ardlethan, 31 km SE (34.37S 147.01 E), 
10.ii.92, CJB (UQIC). Bald Rock Nat. Pk., 30.xi.81, 
M.A. Schneider & G. Daniels (UQIC). Barrington 
Tops S.F., Dilgry River (31.53S 151.32E), 15-16.xi.81, 
T. Weir; Gloucester R (32.04S 151. 41E), 12-14.xi.81, T. 
Weir & A. Calder; Moppy Lookout (31.54S 151. 33E), 
18.xi.81, T. Weir; 3km W Moppy Lookout (31.54S 
151.31E), 18.xi.81, T. Weir & A. Calder; Polbius 
Swamp, 17.xi.81, T. Weir; Thunderbolt's Lkt (31.55S 
151 .30E), 18.xi.81, T. Weir & A. Calder. Batemans Bay, 
4 N, 21.X.52, E.F. Riek. Bathurst, l.v-21.v.62, R.D. 
Hughes; 23.ii.66, M.I. Nikitin (ASCU); 23.X.67 
(ASCU); 26.X.68, N.C. Lloyd (ASCU). Billabong Ck nr. 
Conargo (35.17S 145. HE), 12-17.iv.78, JCC. Boolijah 
Ck via Sassafras, 22.xi.79, JCC. Braidwood, 13 km 
NNW (35.21S 149.44E), 5.xi.81, M.S. Upton. Brooklyn, 
31.X.14 (QMBA). Broken Hill, 100 km SE (32.51 S 
141.37E), 3-13.X.88, E.D. Edwards. Brown Mt., 15.i.69, 
JCC & S.R. Curtis. Bungendore, 13 km E (35.15 S 
149.35E), 6.xi.81, M.S. Upton. Cabbage Tree Ck, Clyde 
Mt., 22.ii.65, D.H. Colless. Cabramatta, l.i.63, M. Ni- 
kitin (QMBA). Condobolin, 17.X.00 (ASCU). Congo, 8 
km SE Moruya, 15.ii.82, M.S. Upton. Coonabarabran, 
51 km N (30.50S 149.26E), 13.11.92, CJB (UQIC). Dai- 
ners Gap (36.12S 148.43E), 1585m, 19.xii.73, 28.xii.73, 
6.ii.74, 20.iii.74, ex. Eucalyptus pauciflora, stcUulata and 



perriuiana forest. Euroka, 4 km W Kempsey, 6.xii.78, 
A. Postle. Forbes, 12.xi.64, D.H. Collass. Fowlers Gap 
Res. Stn (31. OSS 141. 42E), 29.xi-2.xii.81, 8-9.xii.82, 
IDN & JCC. Gibraltar Ra., 15.i.79, rainforest margin, 
IDN. Grafton, 5.iv.36, Dicochrosis puuctiforalis (ASCU). 
Halfway House, Putty Rd., 20.xii.73, IDN (UQIC). 
Hunter R, 3.6 km on Glennies Ck Railway Rd, 12- 
13.xii.78, A. Postle & C. Brennan. Klora nr. Moruya, 
6.iii.66, Z. Liepa. Kosciusko, Sawpit Ck, ll.xii.60, 
D.H. Colless. Lake George nr. Collector, 23.x. 76, Z. 
Boucek (QMBA, USNM). Lansdowne, 3 km N, xi.91, 
riparian rainforest, blossom Watcrhousia floribunda, G. 
& T. Williams. Leeton, 2.ii.66, M.I Nikitin (ASCU). 
London Falls, 12.xii.48, C.E. Chadwick (ASCU). Mac- 
quarie Pass, 7 km ENE Robertson (34.34S I50.40E), 
8.ii.84, IDN. Monga State Forest, (35.38S 149.54E), 
3.xii.77, IDN; 18.ii.83, IDN & JCC. Monga, 5 km SW, 
9.xi.81. Mootwingee Nat. Pk, Homestead Gorge 
(31.17S 142.18E), 7-13. X.88, E.D. Edwards. Mt. Drom- 
edary nr. Narooma, 2100 ft, 4.ii.69, Upton, Taylor & 
Cardale. Mt. Kaputar Nat. Pk., 2.xii.76, E.M. Exley; 
Dawsons Spring (30.17S 150.10E), 1420m, 5-ll.xii.87, 
moericke trap under flowering Kuuzca auilngua, G.R. 
Brown (ASCU). Myalia Tank, 49 km NE Broken Hill 
(31.50S 141.57E), 3.xii.81, IDN & JCC. Narrabri E.F., 
19.ix.60, M.I. Nikitin (ASCU). New England NP, 
Point Lookout, 22.i.79, temperate rainforest, IDN & 
JCC. Orange, 22.ii.66, 23.ii.66, 14.iii.66, M.I. Nikitin 
(ASCU). Orchard Hills, W. Sydney, 26.V.82, K. Helm. 
Parkes, 13.xi.64, D.H. Colless. Pigeon House Ra. via 
Nerriga, 25.x, 22.xi.79, IDN & JCC. Scaly Lookout, nr. 
Coffs Harbour, 6.ix.83, G. Daniels & M. Schneider 
(UQIC). Terrace, 13.xii.78, A. Postle. Tooloom Plateau 
via Urbenville, 10.xi.74, IDN (UQIC). Triangle, 150- 
200m, 5-7.X.79, aerial netting, R. Farrow; Research 
Station, 4.xi.79, aerial net. Wambool Common, 18 km 
ESE Bathurst, 4.iv.80, JCC. West Wyalong, 17 km S 
(34.05S 147.08E), 10.ii.92, CJB (UQIC). Yanco, 4.ii.60, 
M.I. Nikitin (ASCU). Northern Territory: Roe Creek, 
11 km SW Alice Sprmgs (23.46S 133.47E), 9.X.78, JCC. 
Queensland: Applethorpe, 15.x73, J. Rhodes (QDPI). 
Bald Mt. area via Emu Vale, 3500^000', 27-31.1.72, 
S.R. Monteith. Biloela, 25 km E, 13.iii.76, E.M. Exley, 
on Ironbark (UQIC). Boonah, 29.xii.91, 17.vi.92, 
18.vi.92 (UQIC); 16 km N (27.54S 152.41E), 18.ix.94, 
18-19.V.96, 6-7.ix.97, CJB (QMBA). Brisbane, 22.iii.40, 
C.F. Ashby (QDPI); 19.iv.52, S. Barker (UQIC); 
5.xii.77, Eucalyptus, K. Walker (UQIC); iv.78, ex. syr- 
phid larva, B. Cantrell (QDPI); Acacia Ridge, 26.xii.76 
(QMBA); Taringa (27.30S 152.58E), coll. ii.98, em. 12, 
13.iii.98, ex. pupa Dideopsis ac;^rota on citrus, CJB 
(QMBA). Bunya Mts. Nat. Pk. nr. Westcott Plain 
(26.52S 151.34E), 6-7.X.84, IDN & JCC. Charleville, nr, 
13.iv.89, P. Johnson (QMBA). Chinchilla, 6 km W, 5- 
15.iii.98, G. Lithgow (QMBA). Cooloola Nat. Pk., E 
Gympie, 18.X.78, I.D. Galloway (QDPI). Dayboro, 8.5 
km SSE, Sampsonvale cemetery (27.16S 152.52), 
3.ix.95, 12.X.1997, CJB (QMBA). Eidsvold (25.22S 



42 



Journal of Hymenoptera Research 



151.07E), 11.X.84, IDN & JCC. Forest Hill, 18-19. xi. 76, 
M. Tichon (UQIC). Gatton, ZS.iii.SO, P. Twine (QDPI); 
11.V.81, ll.xi.81 (QDPI); D.P.I. Research Stn., 9- 
16.iii.81, 21-27.iv.81, 25.v-l.vi.81, l-7.ix.81, 14- 
21.ix.81, 7-14.ix.81, 14-21. ix.81, 28.ix-5.x.81, 21.X.81 
(QDPI). Gordonvale, ix.20, ex. puparia of cloudy- 
winged syrphid, A.P. Dodd (QDPI). joalah Nat. Pk., 
Tamborine Mt. (27.56S 153.12E), 18-21.X.78, Lawrence 
& Weir. Lemington Nat. Pk, Mt. Bithongabel, 1400m 
(28.16S 153.10E), 23.X.78, Berlese moss & litter Notli- 
ofngus moorei, Lawrence & Weir. Mitchell, bank of 
Mitchell Riv., 9.X.74, I.D. Galloway (QDPI). Monto, 14 
km NW, 12.iii.76, on Eucalyptus, E.N. Exley (UQIC). 
Mt. Beerwah via Glasshouse, 1800', 5.xii.65, T. Weir 
(UQIC). Mt. Glorious, 31.xii.79, IDN; 19-26.xi.79, 24- 
31.xii.79 (QDPI); 22.iii.79, 7.xii.81, E.G. Dahms 
(QMBA); 3.1.82, B. Cantrell (QDPI); 22.vi-18.x.82, 
27.iv.89, 27.iv-26.x.89, 26.x-3.xii, 89, l.ix-17.x.90, A. 
Hiller (QMBA); 17.X.90, E.G. Dahms & G. Sarnes 
(QMBA). Mt. Inkerman (19.45S 147.30E), 28.iv.97, GJB 
(QMBA). Mt. Nebo, xii.61, E. Warwick. Mt. Norman 
area via Wallangarra, 7-8.X.72, S.R. Monteith. Mt. 
Spurgeon, 2 km SSE (16.27S 145.12E), 1100m, 19- 
22.xi.97, GJB (QMBA). Mt. Superbus summit (28.1 4S 
152.23E), 1270m, GJB (QMBA). Mt. Tamborine, x- 
xi.78, Sankowsky (QDPI); 3.iii.84, I.D. Galloway 
(QMBA). Ormiston, iii.6l, ex. aphids, B.R. Champ 
(QDPI). Quilpie, 149 km E (26.33S 145.38E), 20.ix.90, 
M.P. Zalucki & G.V. Maynard (UQIC). Rathdowney 
(2nd Palen Ck. crossing from), 22.iii.75, I.D. Galloway 
(QDPI). Repulse Ck, 23 km NE Bauhinla Downs 
(24.24S 149.23E), 22-23.iv.81, IDN. Reyford, 26.V.78, 
E. Sinclair (QDPI). St. Lucia, University of Queens- 
land, 12.xii.93, ex. pupa of Syrphidae on Souchus sp., 
S.G. Evans (QMBA); 17.xii.95, adult ovipositing into 
pupa Episip-phus viridaurcus, S.G. Evans (QMBA); 
17.xii.95, ex. Episyrphus sp. pupa on Sowhus olcraccus, 
S. Evans (QMBA). Stanthorpe, 12 km SE, 3-30.xii.90, 
3.iv-9.vi.91, G. Sarnes (QMBA); 47 km N, 9.xii.80, on 
Eucalyptus, E.M. Exley & J. King (UQIC). Taroom Dis- 
trict (25.27S 150.03E), Boggomoss 21, ll.xi.96, GJB & 
S. Evans (QMBA). Thornlands, l.xi.80, J.F. Donaldson 
(QDPI). Toowoomba, 28.ii.78, 29.ii.78, ex. Syrphidae 
pupa, B.A. Franzmann (QDPI). Whiskers, 7 km 
WNW Hoskistown (35.24S 149.23E), 29.xi.92, M.S. 
Upton. Wilson's Peak (nr), via Teviot Gap, 700-830m, 
9.i.77, IDN (UQIC). Yatala, 24.xi-23.xii.81, among sug- 
arcane, L.N. Robertson. Yerongpilly, 1-10. i.82, B. 
Cantrell (QMBA). South Australia: Adelaide, reared 
ex. Melaijgyiui viridiccps (Macq.), M. Carver; 50 km S, 
Aldinga Scrub, 5-6.xii.86, JSN. Brookfield Cons. Pk. 
(34.21S 139.29E), 24-26.xi.92, IDN & JCC; SW corner, 
stop 29 (34.24S 139.26E), 20.X.92, Rentz, Roach & 1 lar- 
wood. Cowell, 32 km NNE (32.26S 137.03E), 28.xi.92, 
IDN & JCC; 43 km NNE (33.20S I37.06E), IDN & JCC. 
nr. Lake Eyre South (29.31S 137. 16E), JCC. Lake 
Tungketta (33.46S 135.06E), 30.xi.92, IDN & JCC. 
Lock, 24 km NW (33.32S 135.30E), 30.xi.92, flowers 



Eucalyptus, IDN & JCC. Mernmerna, 33 km N Hawk- 
er (31.36S 138.23E), 17.ix.78, JCC. nr. Moonabbie 
Range (33.17S 137.10E), IDN & JCC. Oraparainna Ck, 
Dingley Dell Camp (31.21S 138.42E), 4-10, 7.xi.87, 
IDN & JCC. Orroroo (32.44S 138.37E), ll.xi.87, IDN 
& JCC. Parachilna Ck (31.08S 138.33E), 8.xi.87, IDN 
& JCC. Parra Wirra Rec. Pk, 30 km NE Adelaide, 
9.xii.86, JSN. Penong, 10 km WNW (31.33S 132.54E), 
14.X.81, IDN & JCC. nr. Pine Hill (33.22S 137.03E), 
28.xi.92, IDN & JCC. Pinnaroo, 18 km SSW (35.25S 
140.49E), 20.X.83, 24.X.83, IDN & JCC; 25 km SSW 
(33.28S 140.47E), 20.X.83, 24.X.83, IDN & JCC; 49 km 
SW (33.42S 140.49E), 20, 24.X.83, IDN & JCC. Port Lin- 
coln, 4 km SW (34.45S 135.49E), 29.xi.92, IDN & JCC. 
Willmington, 2 km SSE (32.39S 138.06E), ll.xi.87, IDN 
& JCC. Tasmania: Bronte Lagoon, 13.1.84, L. Masner 
(CNCI). Bronte Pk., 12 km NNE (42.02S 146.33E), 
2.ii.83, IDN & JCC. Buckland, 5 km W (42.37S 
147.39E), 27.1.83, IDN & JCC. Claytons, Bathurst Har- 
bour (43.22S 146.08E) 13.i.91, Nielson & Edwards. 
Condominium Ck, 3 km WSW Anne (42.58S 146.22E), 
ll.xii.81, IDN & JCC. Cranbrook, 14 km ESE (42.04S 
148.13E), 28.i.83, IDN & JCC. Denson rivulet, N of 
Bicheno (41.48S 148.15E), 6.ii.92, GJB (UQIC). Der- 
went Bridge, 9 km WSW (42.10S 146.08E), 21.1.83, 
IDN & JCC; 18 km SW (42.13S 146.02E), 22.1.83, IDN 

6 JCC. Elephant Pass (41.38S 148.13E), 28.1.83, IDN & 
JCC. Fentonbury, 1 km W (42.39S 146.43E), 12.xii.81 
Franklin R. (42.13S 146.01E), 2.ii.83, IDN & JCC. Frod- 
shams Pass (42.49S 146.23E), 24-23.1.83, IDN & JCC; 

7 km S (42.53S 146.22E), 25.1.83, IDN & JCC; 5 km SW 
(42.50S 146.19E), 24.i.83, IDN & JCC; 8 km SW (42.49S 
146. 18E), 24.1.83, IDN & JCC. Hellyer Gorge, 2.11.67, 
E.F. Riek. Herrick, 1 km NE (41.06S 147.53E), 29- 
30.1.83, IDN & JCC. Kingston, 1 km NE, 26.xii.79, JCC. 
Mayfield Beach (42.15s'l48.00E), 6.11.92, GJB (UQIC). 
Miena, 6 km W (41.59S 146.39E), 20.1.83, IDN & JCC. 
Montumana, 3 km SE (40.58S 143.33E), 19.1.83, IDN 
& JCC. Mt. Barrow via Launcestron, 800-1 000m, 
l.xii.76, IDN (UQIC). Mt. Doris (41.52S I46.03E), 
7.ii.90, coniferous heath, IDN. Mt. Mueller, 3 km NW 
(42.46S 146.25E), ll.xii.81, IDN. Mt. Wellington, 
Shoobridge Bend (42.54S 147.15E), 5.ii.83, IDN & JCC. 
Nelson R. (42.06S 145.44E), 22.1.83, IDN & JCC. Nun- 
amara, 10 km ENE (4I.22E 147.24E), 11.1.83, IDN & 
JCC; 8 km NE, Barrow Ck (41.21S 147.22E), IDN & 
JCC; 11 km NE, Mt. Barrow (41.23S 147.25E), 11.1.83, 
IDN & JCC. Oxford, 4 kmW (42.34S 147.50E), 27.i.83, 
IDN & JCC. Pellon Hut, 3 km S Mt. Oakleigh (41.50S 
146. 03E), iii.91, Lcptospcniiuiii scrub and vicinity, IDN; 
30.xi-8.i.91, open forest; 5-10.ii.90, rainforst, IDN. 
Poatina, 9 km SW (41.48S 146.52E), 20.1.83, IDN & 
JCC; Headrace Adit (41.49S 146.54E), 20.1.83, IDN & 
JCC. Scottsdale, 9 km E (41.10S 147.38E), IDN & JCC. 
The Lea (42..36S 147.19E), 3.ii.83, IDN & JCC. Waya- 
tinah, 3 km NE (42.22S 146.29E), 15, 23.1.83, IDN & 
JCC. Weldborough, 4 km SE (41.14S 147.56E), 13.1.83, 
IDN & JCC. Victoria: Acheron Gap, c. 13 km NNE 



Volume 10, Number 1, 2001 



43 



Warburton, 830m, malaise Nothofagus, D. Pollock & 
L. Reichert. Archeron Way via Warburton, 300-480m, 
16.xii.75, IDN (UQIC). Beech Forest, 10 mi E, l.i.67, 
Z. Liepa; via Colac, 6.i.66, T. Weir (UQIC); Coutts Rd., 
480m, ll.xii.75, IDN (UQIC). Belgrave, 25, 26.xii.26, 
A.P. Dodd (QDPI). Bogong Plains, 5-6000 ft, i.28, F.E. 
Wilson (QDPI). Bruthen, 26.ii.80, IDN & JCC; 9 km N 
(37.38S 147.53E), 8.ii.92, CJB (UQIC). Cann Valley 
H'way, 7 km SW N.S.W. border, 25.ii.80, IDN & JCC. 
Dinner Plain, 11 km from Hotham Heights, 27.ii.80, 
IDN & JCC. Hattah, 12 km NW (34.39S 142.14E), 
19.X.83, IDN & JCC. Jim Jack Ck, 12 km WSW Omeo, 
27.ii.8a IDN & JCC. Kiata, 8 km SSW (36.26S 
141. 46E), 23.X.83, IDN & JCC. Kinglake N. Pk. nr. 
Melbourne, 31.i.77, Boucek (USNM). Lake Crosby 
(35.03S 141.44E), 23.X.83, IDN & JCC. Lind Nat. Pk, 
Growler Ck, 26.ii.80, IDN & JCC. Mitre, 11km NE 
(36.38S 141.48E), 22.X.83, IDN & JCC; 12.5 km NNE 
(36.37S 141.49E), 22.X.83, IDN & JCC. Mitta Mitta Ck, 
25 km NNW Omeo, 28.ii.80, IDN & JCC. Mt. Arapiles 
(36.46S 141.50E), 21.X.83, IDN & JCC. Mt. Donna 
Buang, 1250m, 14-17.1.80, Eucnh/ptii:^ woodland, A. 
Newton & M. Thayer (CNCI); via Healesville, 4080 
ft, lO.i.66, T. Weir (UQIC); via Warburton, 1200m, 
8.xii.76, IDN (UQIC). Mt. Sabine via Barramunga, 
580m, ll.xii.75, IDN (UQIC, QDPI). Omeo, 12 km 
NNW, 28.ii.80, IDN & JCC; 18 km NW, 28.ii.80, IDN 
& JCC. Pirita, 13 km S (34.29S 141. 54E), 18.X.83, IDN 
& JCC. Rye, 27.ii.89, ex. Dialccticn sp. A on Cyiio^loi^- 
siiin aiistralc, R. Sheperd. Yapest, 10 km NW (35.41S 
142.02E), 23.X.83, IDN & JCC. Yarrara, 15 km S 
(34.33S 141 .25E), 18.X.83, IDN & JCC. Western Aus- 
tralia: Boranup Karri Forest, 20 km S Margareb River, 
ll-13.xii.90, A.D. Austin (WARI). Cape Arid NP, 
30.xii.86-31.i.87, JSN; Yokinup Bay area, 31.xii.86- 
3.i.87, JSN. Cape Le Grand Nat. Pk. (33.58S 122.07E), 
lO.i.87, ll.i.87, G. & A. Daniels (UQIC). Condingup, 
c. 55 km E Esperance, 31.xii.86, JSN. Dongara, 30 km 
S., 19.xii.86, JSN. Esperance, 4.i.87, JSN. Fitzgerald 
Riv. Nat. Pk., Quaalup area, 6-9.i.87, JSN. John Forest 
NP, c. 25 km E. Perth, 23-27xii.86, 24-28.xii.86, JSN. 
Ludlow (33.37S 115.29E), 2.xi-23.xii, S.J. Curry. Ned- 
lands, 10.iv.41, from syrphid pupa on rose leaf, K.R. 
Norris. Needilup, 29 km NE (33.54S 119.04E), 30.X.84, 
A. A. Calder. Noongar, 2 km SW (31.21S 118.57E), 
IDN & JCC. Pithara, 2 km SSW (30.24S 116.40E), 
26.ix.81, IDN & JCC. Porongorup Nat. Pk., 1.87, JSN. 
Ravensthorpe, 4.i.87, JSN. Stirling Range Nat. Pk., 
1.87, JSN. Swan River, G. Compere (QMBA); Red 
Gum Spring, 23 km ENE Cranbrook, 20-22.xii.90, 
A.D. Austin (WARI). Walpole-Nornalup Nat. Pk., 17- 
21.i.87, JSN; Nornalup, 5 km SE, 17-18.xii.90, A.D. 
Austin (WARI); Nornalup, 2 km W (34.59S 116.48E), 
17.i.93, E.D. Edwards. Yanchep Nat. Pk., c. 50 km N 
Perth, 20.xii.86, JSN; c. 65 km N Perth, 21.xii.86, JSN. 
Yellowdine, 21 km NE (31.17S 119.53E), lO.x.81, IDN 
& JCC. 

Hosts. — Label data indicate Aphis crnc- 



civora (Koch) (Aphididae) and puparia of 
Dideopsis negrotn (Fabricius), Episyrphus 
viridaureiis Wiedemann and Melangyua vir- 
idiceps (Macquart) (Diptera: Syrphidae) as 
hosts of P. emersoni. There is also one 
anomalous record from Dialectica sp. (Lep- 
idoptera: Gracillaridae). 

Remarks. — Pachyueuron emersoni is dis- 
tinguished by a combination of features 
that are given in the key and description. 
I have seen females, most commonly from 
Western Australia, that lack a marginal 
fringe and therefore resemble P. iielsofii. In 
some instances one or more short regions 
of the wing margin retain setae so absence 
may simply be due to abrasion; however, 
either the setae are for some reason more 
readily lost from females from western 
Australia or presence or absence of the se- 
tae is variable for P. eynersoni in western 
Australia. Females without a marginal 
fringe are differentiated from P. nelsoni fe- 
males by their conspicuously longer mar- 
ginal and postmarginal veins {cf. Figs. 24 
and 36), snioother and shinier medial area 
on the propodeum {cf. Figs. 15 and 29), 
and more elongate petiole that in dorsal 
view is uniformly reticulate {cf. Figs. 22 
and 27). All males of P. emersoni that I 
have seen have a marginal fringe but 
those from western Australia often have 
the lower face only inconspicuously de- 
pressed lateral to the supraclypeal area 
(Fig. 17), much less so than for typical 
specimens from eastern and southern 
Australia (Fig. 16). The western Australian 
males are thus more like males of P. rieki, 
but they do not have the setal patterns of 
the forewing basal fold or the scape as de- 
scribed for P. rieki males. Also, even 
though the facial region may be only in- 
distinctly depressed (Fig. 17), there is still 
a noticeable difference in the reticulate 
sculpture compared with that near the 
eye, the cells being smaller and often more 
obliquely oriented in the depressed re- 
gions (Fig. 18). I have also seen rare males 
of P. emersoni from eastern Australia and 
Tasmania that have a single petiolar seta 



44 



Journal of Hymenoptera Research 



projecting from one side, but these males 
have the lower face distinctly depressed 
lateral to the supraclypeal area and the 
basal fold bare. 

Individuals of P. emersoni are morpho- 
logically very similar to those of P. for- 
mosHui Walker (1833) in Europe and P. al- 
butius Walker (1843) in America north of 
Mexico. However, the propodeum is uni- 
formly reticulate in P. formosum females 
and reticulate with a network of oblique, 
irregular carinae in P. nibutius females. Fe- 
males of both species lack the more or less 
W-shaped complex of plicae and costulae 
and the smoother posteromedian region 
characteristic of P. emersoiii females (Fig. 
15). Males of P. formosum and P. nibutius 
also have the lower face essentially evenly 
convex and uniformly reticulate. 

Boucek (1988) previously suggested that 
P. kingslei/i was only a form of P. emersoni 
but did not formally synonymize the 
names. The feniale lectotype is complete, 
but the antennae are mounted on a slide 
under a separate cover slip from the head 
and antennae of the USNM male paralec- 
totype (Dahms 1983). 

Pachyneuron nelsoni Girault 

(Figs. 25-36, 51) 

Pachyneuron nelsoni Girault, 1928[421]: 2. Holo- 
type female (exaniined). Type data: Austra- 
lia: N. Q., Gordonvale [= Nelsoni, Feb. 1920, 
Dodd. Type depository: QMBA, type no. 
T.9324. Sex described: female. 

Pnciiyncuron aeneus Masi, 1929: 229-231. Holo- 
type female. Type data: Libya (North Africa): 
Oasis of Giarabvib, iii.1927. Type depository: 
Museo Civico di Storia Naturale, Genoa. Sex 
described: female. Synonymy by Boucek, 
1988: 442. 

Atrichoyitilus aeneus; Delucchi, 1956: 141-142. 
Change of combination. 

Paclii/neuron acncuin; Boucek, 1965: 16-18. 
Change of combination. 

Pachyneuron nelsoni; Dahms, 1986: 324-325; 
Boucek, 1988: 442. 

Female. — Body dark with variably dis- 
tinct metallic green luster; antenna brown 
except basal half to all of scape yellow; te- 



gula yellow; legs with femora variably 
darkly infuscate except apically yellowish, 
tibiae and tarsi yellowish. Head with clyp- 
eus flat to slightly depressed and apically 
shallowly emarginate (Fig. 25). Flagelluni 
compact-clavate, with 2 anelli (Fig. 32) 
and 6 funicular segments (Fig. 31); funic- 
ular segments quadrate or slightly longer 
than wide basally to slightly transverse 
apically and with adpressed setae (Figs. 
31, 32); longitudinal sensilla extending al- 
most entire length of funicular segments, 
separated from each other by distance 
equal to 1-2 sensillar diameters (Fig. 32). 
Forewing (Figs. 35, 36) without marginal 
fringe; with relatively inconspicuous, 
white, often spicule-like discal setae; dor- 
sally without line of setae differentiating 
apex of basal cell from speculum; ventral- 
ly without line of setae along cubital fold; 
costal cell with inconspicuous white setae 
on ventral surface (Fig. 35b); veins with 
following ratios (;z = 10): smv/mv = 4.33- 
5.00, mv/mvw = 1.64-2.22, pmv/mv = 
1.25-1.78, pmv/stv = 1.04-1.10. Mesono- 
tum with relatively low convex, broad 
scutellum (Fig. 26). Propodeum (Figs. 26, 
29) with posteriorly convergent, carinately 
margined plical ridges and A-shaped to 
inverted Y-shaped carinae differentiating 
a more or less W-shaped anterolateral re- 
gion and a pentagonal posteromedian re- 
gion, with all surfaces similarly coria- 
ceous-reticulate or with pentagonal region 
more distinctly reticulate; spiracle distinct- 
ly oval. Petiole without setae projecting 
from sides (Figs. 27, 28); in dorsal view 
slightly (up to about 1.3 times) longer than 
wide, with often indistinctly differentiat- 
ed, transverse to quadrate, rugose-reticu- 
late body often having median carina or 
some longitudinal carinae (Fig. 27); in ven- 
tral view completely sclerotized with me- 
dian furrow, the body quadrate to slightly 
transverse, finely longitudinally coria- 
ceous and shiny (Fig. 28). 

Male. — Similar to female except as fol- 
lows: body brighter metallic green or blu- 
ish green; antenna almost uniformly yel- 



Volume 10, Number 1, 2001 



45 



lowish or with flagellum light brown; legs 
uniformly bright yellow beyond coxae; 
scape (Fig. 34) thickest basally and tapered 
toward apex, with anterior surface flat to 
slightly concave over at least basal two- 
thirds and in lateral view with variably 
distinct line of setae along both outer and 
inner anterior margins; flagellum filiform; 
funicular segments oblong, middle seg- 
ments at most 1.75 times as long as wide 
and all segments with very sparse longi- 
tudinal sensilla within apical half of each 
segment and with conspicuous, semierect 
setae about as long as width of segment 
(Fig. 33). Forewing with marginal vein up 
to 2.6 times as long as wide and postn"iar- 
ginal vein up to 1.4 tin"ies as long as stig- 
mal vein; sometimes with 1 or 2 short se- 
tae on dorsal surface of basal fold and 
sometimes with a few short, inconspicu- 
ous setae on dorsal surface within basal 
cell. Propodeum often more uniformly re- 
ticulate with fine or indistinct plical and 
A-shaped carinae (Fig. 29). 

Distribution. — Noyes (1998) recorded P. 
nelsoui from the following regions and 
countries: Afrotropical (Libya), Australa- 
sian (Australia), Oriental (India), and Pa- 
learctic (Moldova, Russia, Turkey). The 
species was additionally recorded from 
Yugoslavia and Azerbaijan by Boucek 
(1977: 46), who stated that it is circum- 
mediterranean. Boucek (1988) stated that 
it is widespread in southern Europe, dry 
countries of Africa and south Asia, and es- 
tablished (probably introduced) in Austra- 
lia. 

Australian distribution (Fig. 51) records 
based on label data of examined speci- 
mens are: Australian Capital Territory: Brindabel- 
la Range, Mt. Ginini (35.32S 148.46E), 24.xi.81, IDN. 
Canberra, 23.xii.30, W.K. Hughes. New South Wales: 
Bowning, nr, 9.xii.69, on Eucalyptus, JCC. Fowlers 
Gap Res Stn (31.05S 141 42E), 29.xi-2.xii.81, on £. ca- 
riialdulciisis flowers, JCC; 8-9xii.82, JCC. Leeton, 
3.ii.66, M.I Nikitin (ASCU). Mootwingee Nat. Pk., Old 
Mootwingee Gorge, 5-8.xi.84, G.R. Brown & H.M. 
Holmes (ASCU). Myaila Tank, 49 km NE Broken Hill 
(31.50S 141.57E), 3.xii.81, IDN & JCC. Orange, 22.ii.66, 
M.l. Nikitin (ASCU); Agric. Res. Stn., 18.viii.93, on 



apple blossom, K. Harding & A. Nicholas (ASCU). 
Triangle, 5-7.X.79, aerial netting 150m, 200-300m, R. 
Farrow; i-iii.85, S.G. Martin, ex. lucerne (ASCU); Re- 
search Station, l.xi79, 4.xi.79, aerial net. Wambool 
Common, 19 km ESE Bathurst, 17.iv.81, JCC. North- 
ern Territory: Alice Springs, 20.ix.78, ex. syrphid 
pupa, L. Rodunz; 7 km NW (23.38S 133.52E), 8.xi.79, 
JCC; 10 km NE (23.37S 133.54E), 6.xi.79, IDN; 35 km 
E (23.41S 134.13E), 25.ix.78, JCC; 39 km E (23.41S 
134.15E), 25.ix, 5.X.78, JCC; 40 km E (23.41S 134.16E), 
5.X.78, JCC; 53 km NE (23.35S 134.22E), 6.X.78, JCC; 
56 km SE (24.1 IS 134.01E), 3.X.78, JCC. Ayers Rock, 
195 km E on Lacsiters Highway, 5.xi.92, P. Danger- 
field (WARl). Queensland: Bramston Beech (17.21S 
146.01E), 14.xii.91, CJB (UQIC). Brigalow Develop- 
ment area, Moura, P.D. Rossiter [S. almum, 21.iv.66] 
(QDPI). Chinchilla, 6 km W, 9-17.X.87, G. Lithgow 
(QMBA). Connors River (22.11S 149.03E), 8.V.80, IDN 
& JCC. Eulo, 32 km W (28.09S 144.43E), 28.X.91, G. 
Daniels, on Flimicrsia maculosa (UQIC). Gatton Col- 
lege Cawes, 30.xi.67, ex. syrphid pupa, B. Teakle 
(QDPI). Gordonvale, 20.ii.20, A.P. Dodd (QMBA). 
Holts Ck, 8 km N Musselbrook Camp (18.33S 
138.11E), 20.V.95, IDN. Miles, 28 km S, 23.ix, D.H. 
Colless. Mount Inkerman (19.45S 147.30E), 28.iv.1997, 
CJB (QMBA). Taroom District (25.27S 150.03E), Bog- 
gomoss 21, ll.xi.66, CJB & S. Evans (QMBA). Towns- 
ville, Ross River, Hermit Pk. (19.18S 146.49E), 4.xii.91, 
CJB (UQIC). Warwick, 9 km S, 13.i.81, J. & C.R. King, 
on Aii;,^ophora costata (UQIC). South Australia: Agnes 
Ck, 44 km NW Granite Downs (26.38S 133. 16E), 
21.ix.78, JCC. Aldinga Scrub, 50 km S Adelaide, 5- 
6.xii.86, JSN. Brookfield Cons. Pk. (34.21S 139.28E), 
24xi.92, 26.xi.92, IDN & JCC. Ceduna, 21 km NW 
(31.56S 133.24E), 14.X.81, IDN & JCC; 32 km NW 
(31.56S 133.24E), 14.X.81, IDN & JCC. nr. Coffin Bay 
(34.38S 135.27E), 29.xi.92, IDN & JCC. Cowell, 43 km 
NNE (33.20S 137.06E), 28.xi.92, IDN & JCC. Edwards 
Creek (28.20S 135.50E), 19.ix.78, JCC. Elliston, 1 km 
SE (33.40S 134.54E), 30.xi.92, IDN & JCC. nr. Lake 
Eyre South (29.31S 137.16E), 18.ix.78, JCC. nr. Moon- 
abbie Range (33.17S 137.10E), 28.xi.92, IDN & JCC. 
Nooltana Creek, 13 km NW Hawker (31.47S 138.21E), 
16.ix.78, JCC. Orapariima Ck, Dingly Dell Camp 
(31.21S 138.42E), 7.xi, 4-10.xi.87, IDN & JCC. Parach- 
ilna Ck (31.08S 138.33E), 8.xi.87, IDN & JCC. Penong, 
10 km WNW (31.53S 132.54E), 14.X.81, IDN & JCC. 
Pinnaroo, 18 km SSW (35.25S 140.49E), 20 & 24.xi.83; 
25 km SSW (35.28S 140.47E), 20 & 24.xi.83, IDN & 
JCC; 49 km SW (35.42S 140.49E), 20 & 24.xi.83, IDN 
& JCC. Taylorville, 12 km ESE (34.08S 140.06E), 
12.xi.87, IDN & JCC. William Creek, 27 km SE (29.05S 
136.31E), 19.ix.78, JCC. Wilmington, 2 km SSE (32.39S 
138.06E), ll.xi.87, IDN & JCC. Yorke Peninsula, 
20.ix.81, aerial netting, R.A. Farrow. Tasmania: Frod- 
shams Pass, 1 km S (42.50S 146.22E), ll.xii.81, IDN. 
Victoria: Haltah, 7 km SE (34.50S 142.18E), 19.X.83, 
IDN & JCC; 12 km NW (34.39S 142. 14E), 19.X.83, IDN 



46 



Journal of Hymenoptera Research 




Figs. 25-30. Pacliyiicuio)! luisoui: 25, head, frontal (9); 26, mesosoma, dorsal ($); 27, petiole, dorsal (?); 28, 
petiole, ventral (6); 29, scutellum-propodeum (9); 30, head, frontal (6). Scale bars = (xm. 



Volume 10, Number 1, 2001 



47 




Figs. 31-36. Pachyneuron nelsoni: 31, antenna (9): 31a, entire, 31b, anelli and tunicular segments; 32, basal 
flagellar segments", fl -fl, (9); 33, antenna {6): 33a, entire, 33b, middle funicular segments, fl^-fl^; 34, scape 
(cJ); 35, forewing, SEM of dorsal surface: 35a, entire, 35b, submarginal vein and costal cell (6); 36, forewing, 
photograph (9). Scale bars = [xm. 



48 



Journal of Hymenof'tera Research 



& JCC. Kiata, 8 km SSW (36.26S 141. 46E), 23.X.83, 
IDN & JCC. Lake Crosby (35.03S 141. 44E), 23.X.83, 
IDN & JCC. Mitre, 11 km NE (36.38S 141. 48E), 
22.X.83, IDN & JCC; 12 km NE (36.37S 141.48E), 
22.X.83, IDN & JCC; 12.5 km NNE (36.37S 141. 49E), 
22.X.83, IDN & JCC. Mt. Arapiles (36.46S 141.50E), 
21.X.83, IDN & JCC. Pirita, 13 km. S (34.29S 141.54E), 
18.X.83, IDN & JCC. Princetown, 5 km NW, 27.xi.77, 
J.F. Donaldson (QDPI). Yapest, 10 km NW (35.41S 
142.02E), 23.X.83, IDN & JCC. Yarrara, 15 km S 
(34.33S 141.25E), 18.X.83, IDN & JCC. Western Aus- 
tralia: Cocklebiddy, 23 km ESE (32.08S 126.18E), 
12.X.81, IDN & JCC. Fitzgerald Riv. Nat. Pk., Quaalup 
area, 6-9.i.87, JSN. Geraldton, 31.xii.75, R. Storey & 
E.M. Exley (UQIC). Kalgoorlie, l.xi.47, swept nr. lu- 
cerne. Ludlow (33.37S 115.29E), 4.xi-22.xii.80, S.J. Cur- 
ry. Madura, 11 km E (31.55S 127.09E), 13.X.81, IDN & 
JCC. 'Marun' CALM Site, 8/4 Prince Frederick Har- 
bour (15.00S 125.21E), 6-ll.vi.88, IDN. Mt. Magnet, 
17.xii.86, JSN. Mt. Singleton, 15 km NE (29.21S 
117.20E), 28-29.ix.81, IDN & JCC. Noongar, 2 km SW 
(31.21S 118.57E), 9.X.81, IDN & JCC. Norseman, 47 
km SSW (32.35S 121.34E), 19.ix.81, IDN & JCC. 
Paynes Find, 5 km SW (29.18S 117.39E), 29.ix.81, IDN 
& JCC. Perenjori, 18.xii.86, JSN. Ravensthorpe, 46 km 
W, 4.i.87, JSN. Yancliep N.P, 20-21. xii.86, J.S.N; c. 50 
km N Perth, on Eiicnh/ptus, 20.xii.86, JSN. 

Hosts. — Noyes (1988) gave Syrphidae 
(Diptera) as the hosts of P. iielsoni, but 
without listing any species; Boucek (1977: 
46) Hsted Episyrphus (= Epistrophe) baltea- 
tus (DeGeer) as an example syrphid host. 
Label data also indicate syrphids as the 
hosts of P. nelsoui in Australia, but exact 
species are unknown. 

Remarks. — Individuals of P. nelsoui are 
most similar to those of P. emersoni and P. 
rieki but are distinguished by the lack of a 
marginal fringe (Figs. 35, 36) in combina- 
tion with a comparatively short and thick 
marginal vein and a shorter postmarginal 
vein (Fig. 36). Individuals also differ 
slightly in propodeal sculpture from those 
of P. emersoni and P. rieki, the propodeum 
having a A-shaped or inverted Y-shaped 
median carina delineating a posteromedi- 
an pentagonal region that is similarly or 
even more conspicuously sculptured than 
is the basolateral W-shaped region (Figs. 
26, 29). Individuals of P. emersoni and P. 
rieki usually have the posteromedian re- 
gion more broadly Pl-shaped, shiny, and 
almost smooth (Figs. 15, 39, 40). The pet- 



iole (Fig. 27) is also shorter than in P. emer- 
soni (Fig. 22) or P. rieki (Fig. 41), but be- 
cause of its length it is often mostly con- 
cealed by the base of the gaster. Antennal 
features further differentiate males of P. 
nelsoni from those of P. emersoni and P. rie- 
ki, the scape having a flat to shallowly 
concave anterior surface that is broad ba- 
sally and tapered apically (Fig. 34), and 
the flagellar segments being comparative- 
ly short (Fig. 33) and usually similarly 
light-colored as the scape. 

The specimen from near Chinchilla, 
Queensland (QMBA) is a gynandro- 
morph, having the head and antennae of 
a male but the metasoma of a female. 

Doganlar (1986) differentiated P. nelsojii 
(as P. aeneum) from other European spe- 
cies of Padnfueuron based on structure of 
the hypopygium and described the new 
species P. erzurumicum, from Turkey, as 
lacking a marginal fringe. He differenti- 
ated the latter species from P. neneum 
based on differences in dimensions of the 
forewing venation and flagellar segments. 
Huang and Liao (1987) also described a 
new species from China, P. nciliatum, as 
lacking a marginal fringe. They compared 
the species with P. grande Thomson but 
did not differentiate it from P. nelsoni, 
though they illustrated a forewing with 
seven setae on the basal fold, three setae 
within the basal cell, and with distinct dis- 
cal setae. 

Pachyneuron rieki Gibson, new species 

(Figs. 37-49) 

Type material. — Holotype, female (ANIC): 
Australian Capital Territory: Flea Ck, 
25.viii.1950, E.F. Riek. Allotype, male 
(ANIC): same data as holotype. Paratypes 
(ANIC, UQIC, CNCI): Australian Capital 
Territory: 7 females, 13 males, same data 
as holotype, the series associated with an 
unidentified syrphid larva (1 female and 2 
males used for SEM). Tasmania: Lake St. 
Clair (42.06S 146.10E), 750m, 25-27.i.l980, 
Lawrence & Weir (1 female). Mt. Doris 
(41.52S 146.03E), 7.ii.l990, coniferous 



Volume 10, Number 1, 2001 



49 



heath, IDN (1 male); 1 km ENE Mt. Ossa 
(41.52S 146.03E), iii.1991, IDN (2 males). 

Etymology. — Named in honour of Edgar 
F. Riek, who reared most of the type se- 
ries. 

Feirmle. — Body dark with metallic green 
luster; antenna dark brown except scape 
yellow; tegula yellow; legs with all except 
apex of femora brown, otherwise yellow- 
ish beyond coxae. Head with clypeus flat 
and apically shallowly emarginate (Fig. 
47). Flagellum compact-clavate, with 2 
anelli (Fig. 44) and 6 funicular segments 
(Fig. 43); funicular segments distinctly lon- 
ger than wide basally to quadrate apically 
and with adpressed setae (Figs. 43, 44); 
longitudinal sensilla extending most of 
length of funicular segments, separated 
from each other by distance equal to about 
2 sensillar diameters (Fig. 44). Forewing 
(Fig. 48) with marginal fringe; with dis- 
tinct discal setae; dorsally with oblique 
line of 7-13 setae on basal fold differenti- 
ating apex of basal cell from speculum 
and with 2-5 setae near apex of basal cell; 
ventrally without line of setae along cu- 
bital fold; costal cell with distinct setae on 
ventral surface; veins with following ra- 
tios {n = 3): smv/mv = 3.33-3.45, mv/ 
mvw = 4.83-5.00, pmv/mv = 1.60-1.76, 
pmv/stv = 1.65-1.76. Mesonotum with 
relatively low convex, broad scutellum 
(Figs. 38, 39). Propodeum (Figs. 39, 40) 
with posteriorly convergent, carinately 
margined plical ridges and less distinct, 
sometimes irregularly n -shaped antero- 
median carina or ridge (costula) near base 
(Fig. 39), the ridges together differentiat- 
ing a more or less W-shaped basal region 
with coriaceously sculptured anterolateral 
depressions from a mostly shiny and 
smooth to finely coriaceous pentagonal or 
hexagonal posteromedian region anterior 
to a coriaceous or medially smooth and 
shiny nucha, with the short region ante- 
rior to Pl-shaped ridge crenulate and the 
surface lateral to plical ridges finely cori- 
aceous (Fig. 40); spiracle distinctly oval. 
Petiole near middle with 1-3 setae pro- 



jecting anterolaterally from each side 
(Figs. 41, 42); in dorsal view about twice 
as long as wide, with distinctly longer 
than wide, uniformly reticulate body (Fig. 
41); in ventral view completely sclerotized 
with median furrow, the body distinctly 
longer than wide, finely longitudinally co- 
riaceous and shiny (Fig. 42). 

Male. — Similar to female except as fol- 
lows: body brighter metallic green or blu- 
ish green; legs uniformly bright yellow be- 
yond coxae; scape entirely yellow, slightly 
expanded basally immediately above rad- 
icle, tapered subapically and slightly 
curved, with line of distinct setae along 
anterior margin (Fig. 46); pedicel some- 
times yellow except brownish dorsally; 
flagellum dark brown, filiform; funicular 
segments elongate, the middle segments 
at least twice as long as wide and all seg- 
ments with very sparse longitudinal sen- 
silla within apical half and with conspic- 
uous, semierect setae about as long as 
width of segment (Fig. 45); forewing with 
basal fold similarly setose as in female but 
sometimes also with 1-3 setae delineating 
posteroapical angle of basal cell and with 
up to 10 setae within cell behind submar- 
ginal vein and toward apex; veins with 
following ratios {fi = 6): smv/mv = 2.70- 
3.23, mv/mvw = 3.84-4.86, pmv/mv = 
1.30-1.74, pmv/stv = 1.52-1.79; petiole 
sometimes without lateral setae (see re- 
marks). 

Hosts. — Unknown species of Syrphidae 
(Diptera). 

Remarks. — This species is most similar to 
P. emersoni, but is distinct based on fea- 
tures used to separate the species in the 
key and descriptions, and as discussed un- 
der the remarks for P. emersoni and P. nel- 
soni. The three males from Tasmania have 
a slightly shorter petiole than the reared 
males from ACT and apparently lack lat- 
eral petiolar setae, though these may have 
been lost during preparation, which in- 
cluded critical-point drying. The three 
Tasmanian males also have somewhat 
shorter submarginal and postmarginal 



50 



Journal of Hymenoptera Research 




Figs. 37-42. Pachyncunvi ncki: 37, mesosoma, dorsal (V ); 38, mesosoma, lateral (9 ); 39, scutellum-propodeum 
( 9 ); 40, apex of scutellum-propodeum, posterolateral {6); 41, petiole, dorsal ( 9 ); 42, petiole, ventral ( 9 ). Scale 
bars = fjLin. 



Volume 10, Number 1, 2001 



51 




43 _2o_ 




Figs. 43^8. Pachyncuron ricki: 43, antenna (9): 43a, entire, 43b, anelli and funicular segments; 44, basal 
flagellar segments, fl,-fl, (9); 45, antemia {6): 45a, entire, 45b, middle funicular segments, fls-fl?; 46, scape 
(S); 47, head, frontal (6); 48, forewing (9). Scale bars = ^x.m. 



52 



Journal of Hymenoptera Research 











-ru\ 




Fig. 49. Austrdlicin distribution: l^dcln/iiciiroii apliidis 
(•), P. nrA-; (A). 

Fig. 50. Australian distribution, I'lnln/nriirou t'liiri- 
so)ii. 

Fig. 51. Australian distribution, Pinln/nmroii //c/sid;; 



veins than do the ACT males (smv/mv = 
2.70-2.86 vs. 2.94-3.23 and pmv/mv = 
1.30-1.43 vs. 1.53-1.74; n = 3), whose ve- 
nation is more similar to that of measured 
females. However, there are insufficient 
specimens of both sexes to accurately es- 
timate true variability in any of the mea- 
sured structures. 

Graham (1969) differentiated P. iDiibra- 
tiim Delucchi (subsequently synonymized 
with P. groenlandicum (Holmgren) by 
Hedqvist, 1977) from P. fonuosum based 
on the presence of 2-12 setae on the basal 
vein. Though this is similar to P. rieki, P. 
groenlandicum lacks the petiolar setae char- 
acteristic of P. rieki and has an evenly re- 
ticulate propodeum. An unidentified spe- 
cies from America north of Mexico has 
petiolar setae and often a setose basal vein 
similar to P. rieki, but differs in propodeal 
sculpture, having the plical region more or 
less evenly reticulate or with some irreg- 
ular, oblique carinae similar to P. albutius. 

CONCLUSIONS 

Without a world species revision it is 
premature to hypothesize about the phy- 
logenetic relationships of the Australian 
fauna of Pachyneuron. However, P. emer- 
sojii, P. nelsoni and P. rieki all share a pos- 
teromedially differentiated propodeal pli- 
cal region that is delineated by a more or 
less W-shaped complex of plicae and cos- 
tulae (Figs. 15, 29, 39). This structure dis- 
tinguishes the species from other morpho- 
logically similar species, such as P. foriiio- 
sujfi and P. albutius from the Nearctic and 
Palearctic regions, respectively, which 
have the propodeal plical region more or 
less uniformly reticulate. Although polar- 
ity is uncertain, the similar propodeal 
structure suggests that P. eniersoni, P. nel- 
soni and P. rieki are closely related and 
may have speciated in Australia, which 
would not support the hypothesis that P. 
nelsofii was introduced into Australia re- 
cently (Boucek 1988). Pachyiwuron fielsoni 
is also one of the most widely distributed 
species in Australia and the only species 



Volume 10, Number 1, 2001 



53 



yet recovered from northern Western Aus- 
tralia. (Fig. 51). The distribution pattern 
does not suggest a recent introduction. Pa- 
chyneuron aphidis is certainly much more 
distantly related to the other species and 
undoubtedly represents a separate intro- 
duction into Australia, probably acciden- 
tally by man into New South Wales based 
on present distribution (Fig. 49) and the 
earliest collection records. 

ACKNOWLEDGMENTS 

This study was conducted primarily during a work 
transfer to the Australian National hisect Collection 
(CSIRO, Canberra), and I gratefully acknowledge 
their assistance and the use of their collection and 
facilities. The individuals and institutions listed un- 
der material and methods are thanked for the loan of 
material on which the study was based. Ms. J. Read 
(ECORC) generated all the illustrations, including 
dissecting and preparing the specimens for SEM and 
photography, producing the images, and preparing 
the illustratory plates. Drs. John Huber and James 
O'Hara are gratefully acknowledgeci for critically re- 
viewing the manuscript. 

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Dahms, E. C. 1983. A checklist of the types of Aus- 
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Dahms, E. C. 1984. A checklist of the types of Aus- 
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Dahms, E. C. 1986. A checklist of the types of Aus- 
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Delucchi, V. 1956(1955). Beitrage zur Kenntnis der 
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De Santis, L. 1957. Anotaciones sobre Calcidoideos 
Argentinos (Hymenoptera). Notas del Museo de La 
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De Santis, L. 1975. Nota sobre calcidoideos neotro- 
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Doganlar, M. 1986. Morphological studies of the hy- 
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Journal of Hymenoptera Research 



Gahan, A. B. 1918. Pwpncln/iiciiron Girault (Hymenop- 
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Gahan, A. B. 1924(1923). Types of two chalcid-flies 
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Gibson, G. A. P. 1997. Chapter 2. Morphology and 
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Girault, A. A. 1916(274]. Australian Hymenoptera 
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Girault, A. A. 1917[322]. The North American species 
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Girault, A. A. 1917(327]. A new genus or subgenus 
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Girault, A. A. 1928(421 (. A prodigeous discourse on ivdd 
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Girault, A. A. 1929(431]. Notes on, and descriptions 
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Hedqvist, K.-J. 1977. Notes on Chalcidoidea XI (Hy- 
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Huang, D. and D. Liao. 1987. A new species of Pa- 



chyneuron (Hymenoptera: Chalcidoidea: Pterom- 
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Kamijo, K. and H. Takada. 1973. Studies on aphid 
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Mani, M. S. 1939. Descriptions of new and records of 
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Mani, m' "s. and G. G. Saraswat. 1974. Part III. Pages 
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of Natural History (2) 5: 125-133. 



J. HYM. RES. 
Vol. 10(1), 2001, pp. 55-75 

New Descriptions of Halictus (Seladonia) from the New World 
(Hymenopterta: Halictidae) 

Jessica Janjic and Laurence Packer 
Department of Biology, York University, 4700 Keele St., Toronto, Ontario, Canada, M3J 1P3 



Abstract. — We describe females of Halictus (Seladonia) pingiiismentus Janjic and Packer ueiv 
species from Guadeloupe Island, Mexico and provide the first descriptions of the male and putative 
queen of H. (S.) harmonius Sandhouse and the male and worker of H. (S.) lauei (Moure). Additon- 
ally, we describe the second known putative queen specimen of H. (S.) lanei and describe the 
huge morphological differences between the castes in this species. Differences among these species 
are discussed with respect to other New World members of the subgenus. 



In order to facilitate the preparation of 
a phylogenetic analysis of bees of the sub- 
genus Seladonia (genus Halictus) with par- 
ticular emphasis on the New World spe- 
cies, we present additional descriptions of 
bees in this group. North American spe- 
cies of the genus Halictus were treated by 
Sandhouse (1941) and Central and South 
American Seladonia by Wille and Michener 
(1971). Hitherto, seven New World Sela- 
donia species have been recognized (Mou- 
re and Hurd 1987): H. (S.) confusus Smith, 
a holarctic species found throughout 
North America and Europe; H. (S.) har- 
monius Sandhouse, apparently restricted 
to southern California; H. (S.) hesperus 
Smith, which is primarily a central Amer- 
ican species, found from Mexico to Co- 
lombia; H. (S.) lanei (Moure), which has 
been recorded from Brazil but which 
seems to extend into Venezuela and Co- 
lombia; H. (S.) lutescens Friese, which is 
found approximately sympatrically with 
H. hesperus; H. (S.) tripartitus Cockerell, 
found in the western USA and northwest- 
ern Mexico; and H. (S.) virgatellus Cock- 
erell, restricted to areas around and above 
the treeline in western North America 
from the North West Territories of Canada 
to New Mexico. 

As a result of our studies we have 



found specimens of an additional species 
collected from the island of Guadeloupe 
off the west coast of Baja California. We 
describe this new species below. We also 
present the first descriptions of the males 
of H. (S.) harmonius and H. (S.) laiiei. Most 
of the aforementioned species of Seladonia 
are known to be social and at least H. hes- 
perus has large morphological caste differ- 
ences (Brooks and Roubik 1983; Packer 
1985) such that the castes would not read- 
ily be recognized as being conspecific. The 
description of H. haniuuuus was apparent- 
ly based upon worker females (some ob- 
servations on sociality in this species will 
be published elsewhere) and that of H. la- 
nei appears, based upon macrocephaly, to 
be that of a queen. Here we provide the 
first detailed descriptions of an apparent 
queen of H. harmonius and worker of H. 
lanei. Lastly, as the original description of 
H. lanei was short and in Portuguese 
(Moure 1940), we provide an additional 
description of a queen of this rare species, 
a specimen which is larger and even more 
macrocephalic than the type. 

We are not undertaking a complete re- 
vision of the New World members of the 
subgenus Seladonia as this is beyond our 
scope at this time. In particular, detailed 
studies of the widespread and variable H. 



56 



Journal of Hymenoptera Research 




Fig. 1. Halictus pinguifiiiwiitus Janjic and Packer, habitus, largest paratype temale. Scale bar = 1mm. 



(S.) coufusus are badly needed and should 
be performed in conjunction with genetic 
studies (Rosenmeier and Packer 1993; Tay- 
lor and Packer 1997). The North American 
Seladoiiia species were keyed and briefly 
described by Sandhouse (1941) and the 
Central American species-pair H. hesperus 
and H. lutcsceiis were treated in detail by 
Wille and Michener (1971). 

MATERIALS AND METHODS 

External morphology is described from 
pinned specimens. Genitalia from male H. 
harrnonius and H. land and the labra of all 
castes /species were removed and treated 
in 5% potassium hydroxide before being 
stored in glycerine. Details of surface 
sculpture were observed with light reflect- 
ed from the light source (a Schott KL 1500- 



Z fibre optic system) using a variety of 
white surfaces, this was found more con- 
venient than using light transmitted 
through semi-opaque paper. Terminology 
generally follows that of Eickwort (1969), 
however, for the labrum we use the ter- 
minology of Walker (1995) and for surface 
sculpture characteristics we refer to 
McGinley (1986). Measurements were 
made using a Leica MS5 microscope with 
an ocular micrometer. When more than 
one individual was available to us we pre- 
sent measurements for the type followed 
by the range in brackets. 

We often refer to lengths of particular 
structures or of pilosity with reference to 
the diameter of the median ocellus "od" 
of the same individual. The relative size 
and density of punctures are given in 



Volume 10, Number 1, 2001 



57 



terms of the relationship between punc- 
ture diameter and the interspaces between 
them such as "i = 2d". Other acronyms 
used are as follows: for metasomal terga 
and sterna we us T and S respectively (as 
the first abdominal segment is the meso- 
somal propodeum this means that T3 rep- 
resents the third metasomal tergum but 
the fourth abdominal tergum), Al refers 
to the first annulus of the antenna (ie fol- 
lowing the pedicel), UID and LID refer to 
the upper and lower interorbital distances 
respectively, lOD is the interocellar dis- 
tance — the distance between the inner 
margins of the lateral ocelli and OOD is 
the ocell-ocular distance, the shortest dis- 
tance between the outer margin of one lat- 
eral ocellus and the ipsilateral compound 
eye. 

In the descriptions below, we concen- 
trate on those features which vary among 
the New World species of Selndonia and do 
not repeat aspects which are constant, or 
almost so, among all 8 species. Character 
states which are diagnostic for a species 
are italicized. 

Halictus (Seladonia) pinguismentus 
Janjic and Packer, new species 

(Figs. 1-8) 

Holotype fe?uale. — Size: Total body 
length 7mm (7-8. 3mm), head width 
2.1mm (2. 1-2. 4mm), forewing length 
5.3mm (5. 3-5. 7mm). Coloration: Head 
brown with dark metallic blue reflections 
except for clypeus, supraclypeal area, and 
epistomal and hypostomal regions non- 
metallic, red-brown; meosoma brown 
with dull metallic green reflections except 
disk of scutum with bronze-metallic re- 
flections, scutellum and metanotum dark 
brown, and legs amber but with tibiae 
slightly darker; metasoma orange-brown 
with apical impressed areas translucent 
testaceous; general body surface quite 
shiny especially on lower face, disk of scu- 
tum and scutellum. Pubescence: Off- 
white, moderately long (1.5-2od), pale or- 
ange recumbent hairs 0.5 od long mixed 



with longer hairs on scutum. Structure: 
Head (Figs. 1-2); Slightly broader than 
long (1:0.9); round in anterior view but 
with vertex flat. Labrum (Figs. 4, 5) with 
basal box comparatively long, length to 
width 1:1.6, apico-medialh/ produced into an 
obtuse angle; elevated median area U- 
shaped; distal process narrow, 7<s"' as wide 
at base as width of basal box, triangular; 
apical keel very broad basally, 0.7 times as 
wide at base as width of distal process at 
that point, gradually tapering to apex in 
dorsal view; keel flat on top, largely semi- 
circular in profile, elongate beyond ventral 
margin of distal process for almost half of 
its length and with this produced portion 
weakly concave ventrally. Mandible (Fig. 
7) long, reaching base of opposing man- 
dible, subapical tooth with dorsal margin con- 
cave, thus apjpearing unusually small at apex 
and expanded basally. Clypeus wide, 3 times 
wider than long, evenly convex, punctures 
uniformly sparse (i > 2od). Supraclypeal 
area with apical margin gently convex, 
punctures of two distinct sizes, apically 
sparser than on clypeus, basally more 
dense than on clypeus. Malar space ex- 
tremely short, approximately as long as 
diameter of ommatidium of compound 
eye. Interocular area with punctures al- 
most contiguous medially, less crowded (i 
= d) laterally and below antennal base. 
Frontal carina extending from just below 
antennal bases to less than half distance 
from its apex to the median ocellus. Eyes 
converging above, UOD: LID 1:0.9. Vertex 
flat and long, 2. Sod, area between lateral 
ocelli slightly raised, lOD = 2od, area be- 
tween lateral ocelli and compound eyes 
flat, OOD = 3.75od; punctures behind 
ocelli crowded, on rest of vertex less 
dense, i<d. Gena long, gradually narrow- 
ing behind eyes in dorsal view, produced 
postero-ventrally to form right angle or 
produced as a rounded lobe; maximum 
width approximately twice maximum 
width of compound eye in lateral view, 
excluding lobe if present; punctures some- 
what effaced in weak striae, striae becom- 



58 



Journal of Hymenoptera Research 




Figs 2-8 HalictHS inn^uisiucutus. 2-3. Head, side view. 2, Largest paratype female. 3, Holotype temaie. 4 5^ 
Labrum. 4. Lateral view. 5, Dorsal view. 6, Pronotum, side view, holotype. 7, Mandible. 8, Femur/ tibial 
junction to show basitibial plate, not to scale. Scale bar = 1 mm. 



Volume 10, Number 1, 2001 



59 



ing stronger ventrally. Hypostomal region 
longitudinally weakly and finely striate, 
concave as a result of the genal angle. Me- 
sosoma: Pronotum (Figs. 1 and 6) with lat- 
eral angle strongly produced, carinate an- 
teriorly, carina continuous with pronotal 
lateral ridge which is strong, acute and en- 
tire; lateral angle concave behind carina in 
dorsal view and then swollen; lateral sur- 
face with one or two additional weak dor- 
so-ventral carinulae; dorsal ridge not car- 
inate. Scutum wider than long (1.2:1); an- 
terior margin evenly convex in dorsal 
view, overhanging pronotum medially; 
median line weak, half length of scutum; 
parapsidal lines weak, extending % length 
of scutum; punctation uniformly deep and 
density, i = 0.5-1 d, moderate in size, be- 
coming effaced along anterior margin of 
scutum. Scutellum 7^=, as long as scutum; 
punctures sparser (i > 2d), shallower and 
smaller than on scutum. Metanotum half 
as long as scutellum; punctures fine, dense 
(i = d), becoming transversely effaced lat- 
erally. Mesepisternum dorso-ventrally 
striate, striations deep and coarse; hypoe- 
pimeral area with striations primarily lon- 
gitudinal. Metepisternum with striae 
which are so deep and coarse as to appear 
more like ridges, primarily directed lon- 
gitudinally. Propodeal dorsal surface in- 
termediate in length between scutellum 
and metanotum; posterior surface carinate 
to M height, sparsely and minutely punc- 
tate; dorsal surface with approximately 30 
longitudinal striae; fine, moderately dense 
punctures at postero-lateral corners; lat- 
eral surface with dorso-ventrally directed, 
weak striae, these absent on anterior por- 
tion which has small, dense (i = d) but 
shallow punctures. Tegula orange-brown, 
shining; with very fine, shallow punctures 
anteriorly. Wings: Veins translucent amber 
and wing membrane hyaline as usual in 
Seladonin. Legs: Hindleg with basitibial 
plate elongate triangular, 2od in length, 
entire (anterior and posterior margins well 
defined) and acutely pointed (Fig. 8); in- 
ner hind tibial spur with 3 or 4 teeth (not 



including apex), the first longer than wide, 
the remainder shorter than basal width. 
Metasoma: Tl length to width ratio 0.7:1; 
length of apical impressed area 3od me- 
dially and 2od laterally; anterior surface 
shining, sparsely punctured (i > 3d), with- 
out background microsculpture; becoming 
weakly, transversely microreticulate at 
brow with punctures minute and dense, 
especially laterally (i > d); microreticula- 
tions absent on disk and punctures in- 
creasingly larger and denser (i = d) pos- 
teriorly, sparser on lateral swellings; api- 
cal impressed region with fine, irregular 
punctures. T2 minutely roughened ante- 
riorly, disk with shallow, dense punctures 
(i = d), apical impression long, 4od, punc- 
tures as described for Tl. Punctures in- 
creasingly small and effaced on successive 
terga, apical impressions of T3 and T4 
long, 4od. Apical hair bands weakly de- 
veloped, not extending ventrally. 

Male. — Unknown. 

Etymology: — The specific epithet literally 
means "fat chin", referring to the expand- 
ed genal region of this species, especially 
in larger specimens. 

Specimens examined. — The holotype fe- 
male is missing both antennae, the left 
mid leg beyond the coxa, and the left hind 
leg beyond the trochanter. The specimen 
appears to have become slightly worn and 
somewhat faded. In addition to the holo- 
type, we designate three paratypes, two of 
which are significantly larger (see below). 
The holotype and one paratype are la- 
beled "Guadeloupe Island, Pac. Ocean" 
(the other two paratypes are labeled 
"Guadeloupe Island P.O."), without date 
or name of collector. Each specimen bears 
a second reddish brown label that has 
"Ent. Soc." typed upon it and all four 
specimens were originally from the Phil- 
adelphia Academy of Sciences Collection 
where all but one are now housed (one 
remains in the Packer collection at York 
University). The two large paratypes have 
several marked differences from the two 
smaller individuals reminiscent of caste 



60 



Journal of Hymenoptera Research 



differences found in some of the other 
New World Seladoiiia species. We describe 
the most important differences below. 

Discussion. — Based upon the appearance 
of the specimens and the labels associated 
with them, they would appear to be quite 
ancient. The locality is an island off the 
west coast of Baja California. It is current- 
ly uninhabited, has been ecologically 
damaged by goats, and is difficult to ac- 
cess owing to steep cliffs on all sides. 

The larger individuals have slight 
bronze reflections on the gena and scutel- 
lum. The gena is produced into a rounded 
lobe postero-ventrally (Fig. 2), this lobe is 
2.5od long and 3od wide at its base and it 
renders the ventral margin of the gena 
concave. The pronotal lateral angle is even 
more strongly developed than in the ho- 
lotype. Additionally, the larger bees have 
4 teeth on the inner hind tibial spur, as 
opposed to the 3 in the holotype. One of 
the larger specimens has much of its sur- 
face covered in an amber coloured mate- 
rial which may be dried nectar, it also has 
much of the pubescence worn away al- 
though its wings are not nicked, suggest- 
ing that it was not an old individual but 
rather one that had been badly treated fol- 
lowing capture. The other large paratype 
is in good condition, although it is missing 
apical tarsal segments of both hind legs 
and the left mid leg. The single small par- 
atype has had its head and thorax partial- 
ly crushed. 

We have not been able to locate any ad- 
ditional specimens of this species. In fact, 
we have not been able to find any addi- 
tional collections of bees from the type lo- 
cality. 

Halicttis (Seladottia) harmonius 
Sandhouse 

(Figs. 10-16) 

Haltctiis (Halictiis) Imnmviiiis Sandhouse 1941: 
36, female (USNM). 

Male. — Size: Very small, total length 4- 
5mm, head width 1.05-1. 2mm, forewing 



length 2.8-3mm. Coloration: Head metallic 
bluish-green except for clypeus, antennae 
and hypostoma; clypeus dark brown with 
apical Ks"' often yellow; antennae reddish- 
brown with scape and pedicel darker and 
anterior surface of flagellum slightly paler 
than remainder, anterior surface of first 
annulus yellowish; hypostoma brown; 
mesosoma nietallic bluish-green except for 
reddish brown venter and legs; legs with 
narrow basal and apical bands on tibiae 
and stripe of varying width on outer sur- 
face of fore tibia, this and mid and hind 
tarsi pale brown. Pubescence: White, 
mostly of moderate length, approximately 
lod; longer (1.5-2od) on face, gena, ante- 
rior of scutum, on scutellum and metan- 
otum, laterally on T5, and apical half of 
T6; short (0.5od) scale-like pubescence on 
side of face, sparse on gena and on apical 
impressions of abdominal terga. Structure 
(Fig 14.): Head: As wide as long but ap- 
pearing longer due to narrowing of clyp- 
eus and of vertex behind compound eyes. 
Labrum wider than long (2.6:1). Malar 
area very short, less than 0.2od. Clypeus 
1.3 times as wide as long; apical % pro- 
jecting below a lower tangent of com- 
pound eye; punctures shallow and sparse, 
i = 3d. Supraclypeal area with punctures 
deeper and denser than clypeus, i = 2d. 
Gena and vertex unmodified. Antenna 
long, reaching past base of metasoma; 
scape short, twice apical width and no 
longer than medial flagellar annuli; pedi- 
cel shorter than wide; Al less than half 
length of succeeding annuli, length and 
width subequal; reniaining annuli twice as 
long as wide or longer. Mesosoma: Prono- 
tum with lateral ridge weakly carinate; 
pronotal angle obtuse; dorso-ventral cari- 
na undefined. Scutum length and width 
subequal; convex anteriorly, slightly over- 
hanging pronotum medially; median fur- 
row weak but faintly discernible for entire 
length of scutum; parapsidal lines distinct, 
extending anteriorly for % length of scu- 
tum; punctation deep, distinct and uni- 
formly dense with interspaces approxi- 



Volume 10, Number 1, 2001 



61 



mately equal to puncture diameters. Scu- 
tellum slightly shorter than 1/2 scutal 
length; median line defined on anterior 
half; punctures uniform as on scutum. Me- 
tanotum half length of scutellum; raised 
medially; punctures dense to rugose me- 
dially, sparse but well defined laterally. 
Mesepisternum minutely roughened, 
punctures effaced. Metepisternum with 
uneven longitudinal striae. Propodeal dor- 
sal surface intermediate in length between 
scutellum and metanotum, 2od long; 
rounded posteriorly in dorsal view; pos- 
terior carinae undefined; dorsal surface 
longitudinally ruguloso-striate, striations 
not reaching posterior margin and finely 
rugose apico-medially; posterior margin 
shining and devoid of sculpture; lateral 
and posterior surfaces with fine punctures 
i > d laterally, sparser on shinier back- 
ground posteriorly. Wings: Veins dark am- 
ber, membrane hyaline. Metasoma: Terga 
slightly sinuate in lateral view (especially 
for T2 and T3), weakly depressed apically 
then gently convex to short (lod) apical 
impressed areas. Tl length subequal to 
width; basal area impunctate and shining; 
punctures deep, fine and dense (i = l-2d) 
elsewhere; punctures increasingly fine and 
sparse on succeeding terga, apical im- 
pressed areas impunctate and shining be- 
neath hair bands. Sterna with short, dense 
pubescence. Apical margin of S4 broadly 
concave, with apically directed tuft of 
hairs on lateral Vi of posterior margin, 
these hairs twice as long as elsewhere, 
hairs medial to these tufts laterally direct- 
ed, short and dense. Apical margin of S5 
almost straight, with marginal row of 
sparse, posteriorly directed hairs, increas- 
ing in length from centre to lateral mar- 
gins of segment. S6 with faint medial, bas- 
al depression. Terminalia (Figs. 15, 16): S7 
triangular with pointed apex. S8 rounded. 
Gonobase with dorso-median suture dis- 
tinct; ventro-lateral margins converging 
posteriorly to very acute lateral projec- 
tions, no medio-dorsal cleft to apical mar- 
gin, dorsally convex in lateral view; gon- 



ocoxae elongate, sides subparallel with no 
marked concavities, strongly reflexed in- 
ner dorsal basal margin, lacking striation. 
Gonostylus enormous, swollen, almost as long 
as gonocoxae, with ventral margin broadly 
rounded and becoming vertical apically, with 
apical lobe glabrous and quadrate both from 
above and in profile, with medial semicir- 
cular concavity half way along length just 
ventral to inner setose lobe. Second gon- 
ostylus half as long as apical gonostylus, 
parallel sided. 

Putative Queen. — Size: Small, body length 
4.3mm, head width 1.3mm, forewing 3mm. 
Coloration: Head metallic green with 
bronze reflections; clypeus and epistomal 
region dark red-brown; gena and hypo- 
stomal area brown with greenish-bronze 
reflections; scutum and scutellum metallic 
green; metanotum dark brown with weak 
green reflections; pleura and propodeum 
metallic blue-green; area between fore and 
mid coxae orange-brown; legs brown, tib- 
iae with pale basal spots; tegula amber; 
antennae brown, paler on anterior surface 
and at apex; metasoma brown; entire body 
except pleura, propodeum, and metasoma 
strongly shining owing to absence of mi- 
crosculpture. Pubescence: Sparse, fine, 
and long (1.5-2od) on clypeus, anterolat- 
erally on scutum, laterally on scutellum 
and metanotum, on pleura, dorsally on 
lateral surfaces of propodeum and later- 
ally on metasomal terga, more abundant 
on T5 and covering T6; short (0.5od) and 
appressed near compound eyes on frons 
and gena, on pronotal angles, and meta- 
notum; apical tergal hair bands weak and 
extending ventrally. Structure: Head (Fig. 
12): quadrate, very slightly longer than 
wide 1.03:1, vertex slightly swollen behind 
compound eyes. Labrum (Fig. 10) with 
basal box twice as wide as long, apical 
margin convex; glandular area less than 
half width of basal box, not strongly pro- 
tuberant, with basal portion lacking glan- 
dular openings medially to give whole 
area a flattened U-shape; distal process an 
equilateral triangle with 17 lateral setae. 



62 



Journal of Hymenoptera Research 



somewhat sparser on apical half; apical 
keel long, extending Vs of its length be- 
yond apical margin of distal process, 
slightly concave ventrally, rounded api- 
cally and somewhat unevenly convex dor- 
sally in lateral view, dorsal surface narrow 
and flat. Clypeus weakly produced medi- 
ally, lateral teeth obtuse; length to width 
1:3.5; punctures small, shallow and un- 
evenly sparse, i = l-5d. Supraclypeal area 
weakly convex apically, punctures as on 
clypeus but somewhat more dense basally 
and laterally, i = l-4d; epistomal lobe 
very obtuse with subantennal sutures 
gradually curving into fronto-clypeal su- 
ture. Eyes converging slightly above (UID: 
LID = 0.95:1). Malar space very short, 
0.25od. Frons with punctures small, dense 
and well defined throughout, i = d, except 
sparser below antennae (i = 1.5-2d) and 
with impunctate area immediately above 
antennal bases; frontal suture short, ex- 
tending from mid-level of toruli to less 
than half distance to median ocellus. Ver- 
tex slightly elongate, 2.5 od from lateral 
ocelli to posterior margin; lOD slightly 
less than Sod; OOD 3. Sod; punctures ef- 
faced behind ocelli. Gena convex without 
angular projections, greatest width to eye 
width ratio slightly less than 2:1; punc- 
tures distinct immediately behind com- 
pound eye (i = d); effaced elsewhere, 
weakly microstriate ventrally; hypostoma 
broadly convex, without angles or projec- 
tions. Mandible (Fig. 11) swollen basally 
with well defined subapical tooth. Meso- 
sonin: Pronotum overhung by scutum me- 
dially; lateral angle obtuse, dorso-ventral 
ridge undefined; lateral carina weak to 
pronotal lobe, collar weakly wrinkled. 
Scutum longer than wide (1.17:1); anterior 
margin straight between pronotal angles; 
median suture broad, extending half 



length of scutum; parapsidial lines weak, 
extending to anterior V3 of scutum; punc- 
tures fine, moderately deep and uniformly 
dense (i = d), but slightly more dense lat- 
erad to parapsidial lines and effaced along 
anterior margin. Scutellum weakly im- 
pressed medially; punctation as on scu- 
tum. Metanotum half length of scutellum; 
punctures minute, dense (i = d) medially 
becoming sparser and effaced laterally. 
Mesepisternum with large, shallow, 
sparse (i = 2d) punctures partly effaced in 
roughened background. Metepisternum 
with antero-posteriorly oriented roughen- 
ing. Propodeum with dorsal face as long 
as scutellum, posterior margin rounded; 
weakly and irregularly striate on dorsal 
surface except rugulose medially; lateral 
surfaces microreticulate, anteriorly with 
weak antero-posteriorly directed striae; 
posterior carinae not developed. Legs: In- 
ner hind tibial spur with 3 teeth, the first 
longer than basal breadth, the other two 
broader than long. Basitibial plate entire, 
acutely pointed, narrow and long (1.7od). 
Wiugs: Veins pale honey coloured, costa 
and prestigma darker, membrane hyaline. 
Mesosoma: Tl length:width 1:1.7; apical 
impressed area lod long; anterior half 
transversely microreticulate; apical half 
with small, weak, and slightly transverse- 
ly effaced punctures of uniform density (i 
= d); apical impressions with exceedingly 
minute, sparse punctures. Punctation in- 
creasingly weak on succeeding terga. Api- 
cal impressed area of T2 lod, of T3 and T4 
longer, almost 2od. Weak apical hair 
bands extending ventrally, that of Tl 
broadly interrupted medially, that of T2 
narrowly so. 

Specimens examined. — We have seen 
males from San Timoteo Canyon, River- 
side Co., California and Yucaipa, San Ber- 



Figs. 9-14. Halictits hmimuiiiis. 9, Labrum, workLM', side view. Lower scale bar, = 1 mm. 10, Labrum, putative 
queen, side view. 11, Mandible, queen. Scale bar = 0.1 mm. 12, Putative queen. Scale bar = 0.1 mm. 13, 
Worker. 14, Habitus, male. Lower scale bar, = 1 mm. 



Volume 10, Number 1, 2001 



63 




64 



Journal of Hymenoptera Research 




Figs. 15-18. Halictiis, terminalia. 15-16. Halictus haiiiuviius. 15, Genitalia, ventral view on left, dorsal on right. 
Scale bar = 0.1 mm. 16, Gonostylus, lateral view. 17-18. Halictus tripartitus. 17, Gonostylus, lateral view. 18, 
Gonostylus, ventral view. Scale bar = 0.25 mm. 



nardino Co., California. The first specimen 
collected was found on August 12"\ 1897 
by H.A. Horn, and is deposited in the Uni- 
versity of California, Riverside collection. 
Two additional males collected in the 
same canyon in 1974 were found in mal- 
aise traps run by M. Wasbauer and R. Mc- 
Master on Sept. 9"', these are in the Cornell 
University collection. The male from Yu- 
caipa was collected by T. Griswold on 
June 9"' 1975 and resides in the USDA col- 
lection at Logan, Utah. The queen speci- 
men was collected at Wildwood Canyon, 
San Bernardino Co., California on May 
22'^^', 1977 by Terry Griswold who recog- 
nised the macrocephalic nature of the 



specimen and labeled it as such. It resides 
in the USDA bee lab, Logan, Utah. 

Discussiofi. — Males of H. harmonius vary 
in the extent of pale colouration on the 
legs and clypeus. Some specimens have 
the clypeus entirely dark and some have 
almost the entire surface of the fore tibia 
pale in colour. All, however, have the mid 
and hind tibiae dark and concolorous with 
the corresponding femur. 

Variation among the females would ap- 
pear to result from features that probably 
relate to caste differences. The large pu- 
tative queen specimen differs from the 
smaller workers that we have seen pri- 
marily in having a large, quadrate head 



Volume 10, Number 1, 2001 



65 



which is slightly longer than wide and 
eyes that are slightly divergent below 
(compare Figs. 12 and 13). In contrast, the 
worker head is slightly wider than long, 
more rounded and the eyes are slightly 
convergent below. The worker also has a 
more strongly produced clypeus; the wid- 
ened lower portion of the face of the pu- 
tative queen giving the clypeus a flatter 
aspect. The labra differ in that the queen 
has a longer and slightly differently 
shaped apical portion and a basal portion 
with a more abrupt apical margin than is 
found in the worker (compare Figs. 9 and 
10 for worker and queen, respectively). 

This species is most readily distin- 
guished from other North American mem- 
bers of the subgenus by its extremely 
small size. Indeed, these bees are small 
even for Lnsioglossum (Dialictus), from 
which they can be readily distinguished 
by the usual Halictus characters of strong 
apical wing veins and apical bands of pu- 
bescence on the abdominal terga (contrast 
figures 143 with 145 and 146 with 147 in 
Michener et al., 1994). Based upon geni- 
talic characters (see below), this species is 
most closely related to H. tripnrtitiis, from 
which it is readily distinguished on the 
basis of size (the smallest H. tripnrtitus are 
fully 50% larger than the largest H. har- 
)iioiiius) and the greater density of punc- 
tation of head, mesosoma and metasoma 
in the larger species. Additionally, fresh 
females of H. tripartitus have very well de- 
veloped snowy-white apical tergal bands 
of pubescence whereas those of H. harfnou- 
ius are sparser and on Tl are very broadly 
interrupted. 

The genitalia of male H. harmoiiius are 
very distinctive and differ markedly from 
all other New World Seladonia except H. 
tripartitus (compare Figs. 15 and 16 with 
18 and 17, respectively). Both species have 
large, swollen, sinuate gonostyli with a 
medial semicircular concavity and a thick, 
glabrous, apical lobe instead of a narrow, 
setose process. These gonostylus charac- 
ters are shared by no other Halictus species 



known to us. The two species differ in that 
H. hartiiouius has a proportionately much 
larger gonostylus — as long as the gono- 
coxae (Fig. 15), whereas this feature is 
only Vi as long as the gonocoxe in H. tri- 
partitus. The shape of the apical lobe of the 
gonostylus also differs markedly between 
the species. In H. harmonius it is quadrate 
in profile (Fig. 16), whereas in H. triparti- 
tus it is dorso-ventrally flattened (Fig. 17). 
The shape of the gonostylus just basal to 
the apical lobe is also different — in H. har- 
monius this area is angularly emarginate 
whereas in H. tripartitus there is a deep, 
sharp cleft that extends to the inner-basal 
margin of the lobe and the area basal to 
the lobe is expanded apically to form a 
short shelf beneath the apical lobe. The 
male of H. tripartitus also has paler legs 
and a head that is slightly wider than 
long, as opposed to round, and is not so 
markedly narrowed behind the com- 
pound eyes as in H. harmonius. 

Halictus harmonius is a rare species hav- 
ing been found most often in the region 
around Yucaipa and the San Timoteo can- 
yon in the San Bernardino /Riverside re- 
gion of California. We have been unable 
to verify the records of this species listed 
in Krombein et al. (1979) from Colorado. 
We presume that their records were erro- 
neous as this state was not included in the 
list of localities given by Moure and Hurd 
(1987). 

Halictus (Seladonia) land (Moure) 

(Figs 19-23, 26-29, 32-36) 

Pachycehle land Moure 1940:55. 
Halictus (Seladonia) land Michener 1954:38 
(Moure Collection). 

Queen. — Size: Total body length 9.5mm; 
head width 3.1mm, forewing length 
6.8mm. Coloration: Head red-brown on 
clypeus, supraclypeal area, genae, and hy- 
postomal area, with bronze-green metallic 
reflections elsewhere; malar area and apex 
of mandible black; antennae light brown. 
Scutum and scutellum dark brown with 



66 



Journal of Hymenoptera Research 





Fig. 19. Halktus hvm, macrocephalic female head FigS- 20-22. Halicius lauei labra. 20-21. Queen. 20, 
(center) with worker (lower left) to show size differ- Dorsal view. 21, Lateral view. 22, Worker, lateral 
ence between castes. view. Scale bar = 1 mm. 



bronze-green reflections anteriorly on scu- 
tum and medially and along lateral mar- 
gins of scutellum; remainder of mesosoma 
greenish-blue metallic except legs light 
amber-brown. Metasomal terga light 
brown on Tl-3, T4 and T5 almost black, 
with weak metallic green reflections 
throughout, though somewhat more 
strongly in more posterior segments. En- 
tire head and dorsal surface of mesosoma 
very shiny, completely lacking in back- 
ground microsculpture. Pubescence: Off- 
white except for short appressed pubes- 
cence on metasomal terga 3 to 5 and all 
pilosity on the legs which are golden; 
mostly 1 od in length and sparse, partic- 
ularly on dorsum of mesosoma. Structure: 
Head (Fig. 19): Massive, avisiderabli/ wider 
than thorax, ratio of head widtli to iiitertegular 
distance 1.6:1; shape quadrate, slightly 



wider than long. Labrum (Figs. 20, 21) 
with basal box only 1.5 times as wide as 
long, parallel sided, apical margin medi- 
ally produced to give a pronounced V 
shape but with apex somewhat rounded; 
glandular area very weakly produced, U 
shaped, pores sparse; distal process elon- 
gate, almost 1.5 times as lojig as basal width, 
laterally weakly convex, apex broadly rounded; 
marginal setae becoming widely separated in 
apical half where they arise from the dorsal 
surface of the labrum rather than the lateral 
)mirgins; median keel bisinuate, apical half 
with dorsal margin transversely concave, 
apical margin of keel concave; ventral sur- 
face of labrum completely flat. Mandibles 
enormous, reaching inner ventral margin 
of contralateral compound eye, narrowed 
and slightly outwardly curved beyond 
subapical tooth. Clypeus wide and short, 
length to width 1:4; very weakly convex. 



Volume 10, Number I, 2001 



67 









Figs. 23-31. Hnlictus lanci and H. hcsperus. 23, Worker scutum showing punctation of H. laud (left side) and 
H. hesperus (right side). 24-25. H. hcsperus, propodea. 24, Worker, 25, queen. 26-27. H. land, propodea. 26, 
Worker, 27, queen. Scale bar = 0.3 mm. 28-29. H. land, basitibial plate. 28, Worker. 29, queen. 30-31. H. 
hesperus, basitibial plate. 30, Worker, 31, queen. Basitibial plates not to scale. 



68 



Journal of Hymenoptera Research 



zuith blunt median tubercle, slightly de- 
pressed lateral to the tubercle, lateral clyp- 
eal teeth very short and obtuse; dorsal 
margin for the outermost quarter on each 
side completely straight and slightly ori- 
ented anteriorly, epistomal angle obtuse; 
punctures large and shallow with i = d. 
Supraclypeal area triangular, apical width 
twice its length, very weakly convex; 
sparsely punctured i = 2d. Compound 
eyes convergent above, UOD: LID 7:8; un- 
usually small in comparison to reminder 
of head, length only 0.6 that of head. Fron- 
tal carina very short, extending from level 
witli ventral margin of antennal socket to just 
above them. Punctures small and moderate- 
ly dense, most dense (i ^ d) around ocelli, 
becoming larger and sparser both anteri- 
orly and posteriorly, particularly sparse 
on vertex (i = 3d). Vertex swollen, such 
that head slightly wider at some distance 
behind compound eyes than across them; 
very long such that distance betiveen lateral 
ocellus and posterior margin of z'ertex = 6od 
(Fig. 19); OOD 4od, lob 2.5od; ocelli sit- 
uated in shallow depressions. Gena pos- 
tero-ventrally greatly elongate, almost 
twice as long as greatest width of com- 
pound eye, giving head a triangular ap- 
pearance in lateral view. Hypostonia 
broad and flat, without teeth or other pro- 
tuberances; hypostomal carina strong, 
particularly posteriorly. Mesosoma: Prono- 
tum with lateral angles quadrate, stroiigly 
produced beyond scutum both anteriorly and 
dorsally, carinate anteriorly, carina contin- 
uous with strong pronotal lateral ridge; no 
carina on dorsal ridge; lateral surface with 
strong dorso-ventral striae. Scutum wider 
than long, ratio 6:5; median furrow deep such 
that anterior margin of scutum is biconvex, 
extending half length of scutum; parapsi- 
dal lines distinct, extending from near 
posterior margin to anterior y'-ths of scu- 
tum; punctures shallow, small, and 
sparse, i = 1.5d, except near antero-lateral 
corners where i = d. Scutellum 2.5 times 
as wide as long, T^^ths as long as scutum, 
very flat; punctures as on scutum but 



denser around margins. Metanotum half 
as long as scutellum, uniformly, densely, 
and minutely punctured. Mesepisternum 
dorsoventrally striate on lateral and ante- 
rior surfaces, striae continuing transverse- 
ly on ventral surface, posterior margin 
lacking striae, with a few weak punctures, 
this non-striate region is longer ventrally. 
Metepisternum dorsoventrally striate as in 
mesepisternum. Propodeum with dorsal 
surface two-thirds as long as scutellum, 
with approximately 40 fine, longitudinal 
striae which are strongly curved to give an 
almost fi}igerpri}it-like pattern (Fig. 27); stri- 
ae do not reach apical margin of dorsal 
surface medially, space between striae and 
apical margin completely devoid of sculp- 
ture and very shiny; lateral surface micro- 
reticulate with sparse, minute punctures. 
Wings: veins amber except for costa, 
which is darker brown, membrane hya- 
line. Legs: Basitibial plate of hind leg with an- 
terior margin only well defined at the apex, 
posterior margin well defined, apex pointed 
(Fig. 29); hind tibial spur with four teeth, 
the first twice as long as broad, the second 
as long as broad, the third and fourth suc- 
cessively shorter. Metasoma: Anterior of Tl 
microreticulate with small, weak, widely 
spaced punctures (i > 3d); disk with very 
small, dense punctures, i = d; apical im- 
pressed region weakly differentiated from 
remainder of tergum, 2od in length, simi- 
larly punctured as on disk; punctures on 
remaining gastral terga minute and some- 
what sparser than on Tl; apical impressed 
areas poorly differentiated; apical hair 
bands extremely weak, not extending ven- 
trally. 

Worker. — Size: Body length 5.8-6.5mm, 
head width 1.6-1. 8mm, wing length 4.5- 
5mm. Coloration: Metallic green through- 
out except for apical Vs of clypeus dark 
brown; mandible amber with red-brown 
apical 1/3 and dark brown basal spot; an- 
tennae pale brown; femur and posterior 
surface of fore and sometimes mid-tibia 
brown, remainder of tibiae and tarsi am- 
ber; mesosomal venter brown, apical im- 



Volume 10, Number 1, 2001 



69 



pressions of terga translucent. Pubes- 
cence: Hairs white, slightly off-white on 
pleura and abdomen; mostly lod long. 
Structure: Head: Slightly wider than long, 
rounded. Labrum (Fig. 22) with basal box 
twice as wide as long, anterior margin 
convex; glandular area strongly produced, 
medially divided almost completely by 
non-glandular depression to give it a bi- 
convex shape in apical view; distal process 
longer than basal width, laterally gently 
bisinuate, convex apically; with approxi- 
mately 27 lateral setae; apical keel project- 
ing for K:, its length beyond apex of distal 
process, flat ventrally, convex dorsally, 
subtruncate apically; narrowed to sharp 
dorsal margin. Mandible extending only 
slightly beyond opposing clypeal tooth; 
with blunt subapical tooth. Clypeus 
length to width 1:2.5; evenly and quite 
strongly convex; punctures weak, highly 
variable in size, anteriorly open and un- 
even in density with i — d apically and 
basally, i = 2-3d on disk. Supraclypeal 
area with anterior margin only slightly 
convex, almost straight; punctures as on 
clypeus but more clearly bimodal in size, 
with larger ones more numerous. Frontal 
suture extending from below antennal 
sockets to half the distance to median ocel- 
lus. Inner eye margins subparallel UID — 
LID. Frons with punctures larger, shallow- 
er, partly effaced and sparse (i = 1.5d) 
lower on face to smaller, deeper, entire, 
and denser (i = d) below ocelli. Vertex 
with lOD = 2od, OOD = 2od and distance 
from lateral ocelli to posterior margin of 
vertex 2od; punctures becoming increas- 
ingly sparse, small and effaced posterior- 
ly. Gena with greatest width barely any 
greater than that of compound eye (1.1:1); 
without processes or angulation; punc- 
tures weak and effaced. Hypostoma flat 
without processes. Mesosoma: Pronotal lat- 
eral angle obtuse, lateral ridge weak, no 
stronger than the few, more posterior dor- 
so-ventral carinae. Scutum slightly wider 
than long (1.1:1); straight between prono- 
tal angles; slightly overhanging pronotum 



medially; punctures small, shallow, slighth/ 
effaced trausversely and sparse (i = 1.5^d), 
somewhat more dense and even laterad of 
parapsidal lines (Fig. 23). Medial suture 
extending half length of scutum, deeply 
impressed anteriorly; parapsidal lines ex- 
tending from posterior Va of scutum to an- 
terior Vs; ending in small pits both anteri- 
orly and posteriorly. Scutellum medially 
unimpressed; punctures as on scutellum, 
but more uneven in size and density. Me- 
tanotum half length of scutellum; im- 
pressed medially; punctures effaced, sur- 
face shiny. Propodeum with dorsal sur- 
face intermediate in length between scu- 
tellum and metanotum; with striae 
transverse medially on basal half, arcuate, 
open posteriorly on posterior half, trans- 
verse laterally; in its entirety area appears 
like a broad fingerprint (Fig. 24); lateral 
surface with weak, effaced punctures in 
minute dorso-ventral roughening. Mese- 
pimeron with weak, effaced punctures, 
background weakly dorso-ventrally mi- 
crostriate; hypoepimeral area with weak, 
broad striae directed antero-dorsal to pos- 
tero-ventrally. Metepisternum with coarse 
antero-dorsal-postero- ventrally directed 
weak, irregular striae. Wings: Veins pale 
straw in colour, membrane hyaline. Legs: 
Basitibial plate of hind leg with anterior mar- 
gin loeakh/ defined at apex only, posterior mar- 
gin well defined and sinuate (Fig. 28). Inner 
hind tibial spur with 3 broadly rounded 
teeth, first much larger than others. Meta- 
soma: Tl longer than broad (1.2:1); apical 
impressed area much wider medially 
(3od) than laterally (lod); background 
sculpture transversely microreticulate 
with small, weak punctures (i < 1.5d) in 
anterior half, punctures becoming stron- 
ger and background sculpture weaker in 
apical half; apical impression minutely 
and sparsely punctured. T2 with punc- 
tures slightly larger and deeper than Tl, 
without microsculpture. T3 with punc- 
tures more effaced, even more so on T4 
and T5. Apical impressed areas on T2-5 
2od in length. 



70 



Journal of Hymenoptera Research 



Male. — Size: Total body length 7mm, 
maximal head width 1.5mm, forewing 
length 4.7mm. Coloration: Head metallic 
bluish green with slight bronze reflection 
medially, except clypeus which is brown 
with bluish green reflection basally, paling 
to yellowish amber non-metallic on apical 
V3. Scape and pedicel brown, flagellum 
pale brown dorsally, dark amber ventral- 
ly, first annulus amber throughout. Pro- 
notum and mesosomal venter brown with 
metallic bluish green reflection; mesoso- 
mal pleura and terga metallic green; pro- 
podeum metallic bluish green. Legs amber 
with fore and hind coxae pale brown. Me- 
tasomal terga metallic golden green, more 
weakly so on apically impressed areas. 
Sterna dark testaceous. Pubescence: White 
to cream coloured, mostly long (1.5-2o.d.) 
on face, gena, hypostoma, scutuni, scutel- 
lum, metanotum, and mesosomal pleura. 
Shorter (1 o.d.) on vertex, even shorter (% 
o.d.) on anterior interocular area, inter- 
mixed with long, fine, and golden hairs on 
metasomal terga, and creamy white with 
pronounced branching on posterior pro- 
podeum, and laterally and anteriorly on 
metasoma. Very short {Vi o.d.), broadly 
plumose, and appressed on supraclypeal 
area, clypeus, dorsal surface of collar and 
of posterior lobe of pronotum, lateral and 
posterior margins of scutum, metanotal 
anterior margin, and on apical impressed 
areas of metasomal terga in worn or in- 
complete bands. Structure: Head: as wide 
as long, but appearing longer owing to 
narrowing at level of clypeus. Labrum not 
visible in undissected specimen. Malar 
area short, approx. 0.5 o.d. Clypeal width 
subequal to length; apical 2/3 projecting 
below lower tangent of eye margins; 
punctures large, shallow, and dense (d>i), 
apically effaced. Supraclypeal area with 
smaller punctures apically and much 
smaller between toruli, d>i. Intraocular 
punctures small, and longitudinally ef- 
faced between torulus and compound eye; 
small, deep, distinct, and crowded (d>2i) 
on frons; slightly larger on vertex, effaced 



and more widely spaced posteriorly on 
vertex. Gena and vertex unmodified. An- 
tenna reaching posterior margin of scutel- 
lum; scape 3 times as long as apical width; 
pedicel shorter than wide; Al shorter than 
wide, all remaining flagellar annuli twice 
as long as wide. Mesosoma: Pronotum with 
lateral ridge weakly angulate; pronotal an- 
gle wide but distinct; collar chagrined. 
Scutum length % width; slightly produced 
anteriorly between pronotal lateral angles, 
otherwise flat anterior margin; not over- 
hanging pronotum; median furrow dis- 
tinct, extending half length of scutum; par- 
apsidal lines weak, extending to anterior 
VS; punctures moderate in size and deep, 
d>i, weaker and slightly effaced anteri- 
orly, slightly sparser (d = i) posteriorly. 
Scutellum slightly less than half as long as 
scutum; punctures small anteriorly, d = 
2i; becoming denser, larger, and irregular 
posteriorly, where d~i; most dense on 
postero-lateral corners and posterior mar- 
gin. Metanotum less than Vi length of scu- 
tellum; very narrow medial longitudinal 
area marked by very small contiguous 
and deep punctures; lateral to this punc- 
tures small, d = i; extreme lateral margin 
with punctures larger and effaced into ru- 
gae. Mesepisternum rugose to weakly 
dorso-ventrally striate anteriorly; hypoe- 
pimeron rugose. Metepisternum rugoso- 
striate in antero-posterior direction. Pro- 
podeum length subequal to scutellum; ar- 
cuate striate with arcs open posteriorly, 
striae confused medially, giving them im- 
pression of rugae; smooth at postero-lat- 
eral margins; laterally rugose; posterior 
face with small punctures d>i; lateral ca- 
rinae extending nearly half height. Wings: 
Veins, prestigma, and stigma pale honey 
coloured, costa brown, membrane hyaline. 
Metasoma: Terga with apical impressed ar- 
eas up to lo.d. long medially and 0.5 o.d. 
laterally, most distinctly marked on Tl, 
and weaker and shorter on subsequent 
terga to imperceptible as an impression on 
T5; dorsal surface not sinuate in profile. Tl 
60% long as wide; lateral swellings dis- 



Volume 10, Number 1, 2001 



71 




Figs. 32-33. Halicttis sterna. 32, H. land, sternum 5, 
apical view. 33, Sternum 6, ventral, H. Iwsperus (right), 
H, Iivwi (left). 



tinct; area anterior to brow transversely 
microrugulostriate, brow with shining 
background and small punctures (d<i); 
punctures on disc dense, d>2i, sparser 
across lateral swellings; punctures on im- 
pressed area smaller and very dense 
(d>i). Punctures small, d = i, becoming 
increasingly more effaced on subsequent 
terga; apical impressed areas of subse- 
quent terga with punctures as on discs, 
slightly smaller and sparser on T5 and T6. 
S4 weakly concave apically. S5 with a short 
ifiedio-apical tuft of fine erect Jmir, Va o.d. long 
and wide, hairs % o.d. long (Fig. 32). S6 
with wide medial longitudinal impuncta- 
te, glabrous, depressed area bordered by 
long hairs (Fig. 33). Termijuilia: S7 trian- 
gular with pointed apex. S8 with triangular 
medial projection on rounded stepped base 
(Fig. 34). Gonobase long, concave lateralh/; 
median suture distinct only basally; small 
medio-dorsal cleft in apical margin; ante- 
rior ventral margin obtusely angled (Fig. 
35); ventro-lateral arms converging apical- 
ly, though not touching. Gonocoxae with 



latero-basal concavity; ventral bridge very 
deep, basal margin deeply and abruptyl 
concave; lacking dorsal striations. Gonos- 
tylus with body approximately half length 
of gonocoxe; rectangular in lateral view 
with basal width slightly shorter than api- 
cal width and with dorsobasal margin pro- 
duced; apical inner surface bearing numer- 
ous hairs; inner margin bearing 6 thick 
and apically swollen hairs directed mesad; 
anteriorly directed portion of recurved apical 
projection narrowed to a blunt point; apical 
stylus long, % length of gonostylus body, 
narrow, recurved dorsally, and bearing 
long branched hairs. Second gonostylus 
absent. Penis valve with tips dorsoven- 
trally flattened, bluntly pointed, and api- 
cally slightly recurved dorsad; penis valve 
ventro-basal projection long and narrow. 

Specimens examined. — We have observed 
workers from Conceicao do Araguaia, 
Para, Brazil, collected in July, and from 
Lara, Venezuela, collected in June, and 
from Merida, Venezuela, no date from the 
Cornell University collection. The single 
male was found in a collection at the Car- 
negie Museum of Natural History, bearing 
the label: Boqeirao, Rio Grande, Brazil, 
collected on January 8, 1908, also labeled 
Carn. Mus. Ace. 3533 along with a series 
of 9 females labeled Barra Bahia, Brazil, 
Dec. 6 1907, Carn. Mus. Ace. 3533. 

The large female specimen described 
here has been identified by Padre Moure 
as belonging to Pachyceble lanei, the name 
by which this species was known prior to 
its recognition as a member of the genus 
Halictus by Michener (1954). It bears no lo- 
cality label and is in the Cornell Univer- 
sity collection. Our specimen is larger and 
hence somewhat more macrocephalic than 
the type, photographs of which were 
kindly sent to us by Dr. Danuncia Urban 
of Curitiba. More interestingly, our speci- 
men has the striations of the propodeal 
enclosure partly longitudinal, much more 
so than the type specimen or the workers, 
in which they are primarily transverse. 
While it is not impossible that this single 



72 



Journal of Hymenoptera Research 




Volume 10, Number 1, 2001 



73 



specimen represents a species distinct 
from H. lanei we take the more conserva- 
tive position and ascribe the variation in 
propodeal sculpture to allometric varia- 
tion. In support of this conclusion is the 
observation that queens of the closely re- 
lated H. hesperiis also have most of their 
propodeal striae longitudinal whereas 
those of the workers are transverse (com- 
pare Figs. 24 and 25 for worker and queen, 
respectively). 

Discussion. — Only two other Seladouin 
species come close to the geographic range 
of H. Innei: H. hesperus, and H. lutescens. 
Females of the latter are readily separable 
from those of the other two species be- 
cause of their largely orange metasoma, 
dense punctation on the scutum with i<d 
and their entirely rugulose dorsal propo- 
deal surface. Both queens and workers of 
H. lanei are distinguishable from those of 
H. Jiesperus by the incomplete basitibial 
plate of the hind leg (Figs. 28-31). In H. 
hesperus the basitibial plate is entire 
whereas in H. Imiei it is absent anteriorly 
except at the extreme apex. This reduced 
anterior margin of the plate is also shared 
by both large and small females of H. lu- 
tescens. Furthermore, queens of H. hesperus 
have an angle on the hypostomal carina 
rather than on the gena (Brooks and Rou- 
bik, 1983), the latter being evenly convex 
in both H. hesperus and H. lutescens, but 
markedly produced in large specimens of 
H. lanei as noted above. 

Differentiating between the workers of 
H. hesperus and H. lanei is more problem- 
atic. Other than the basitibial plate char- 
acter mentioned above, the most readily 
detectable difference appears to be in the 
nature of the scutal punctation (Fig. 22). 
In H. hesperus the punctures are shallow 



and largely effaced in an approximately 
triangular area between the central point 
of the scutum and its antero-lateral cor- 
ners. Elsewhere they are better defined, 
separated by interspaces that approximate 
their own diameters and quite variable in 
size with some comparatively large and 
shallow punctures among the rest. In con- 
trast, the scutal punctures of H. lanei are 
partly effaced throughout the scutum, no- 
where are they as strongly effaced as in 
the anterior region of the scutum of H. hes- 
perus but they are more uniformly so. The 
punctures of H. lanei workers are also 
more widely spaced, with i > 1.5od. 

Most of the interesting aspects of the 
morphology of the large specimen — the 
labrum (Figs. 20, 21), the extreme length 
of the mandibles (Fig. 19), enormous swol- 
len head, very long gena, and the enlarged 
pronotal angles — are attributable to ex- 
treme caste dimorphism. Indeed, with a 
head width of 3.1mm, this queen-like in- 
dividual is twice as large as the smallest 
worker we have available for study (Fig. 
19). Translating these linear measure- 
ments into mass, it is possible that this 
queen weighs 8 times as much as the 
smaller workers. Halictines take approxi- 
mately 8 foraging trips to produce a pol- 
len ball that yields an individual of the 
same size as the forager. It is known for 
H. hesperus that the workers produced by 
the queens are smaller than the later 
emerging workers (Packer, 1985). This 
leads to the intriguing possibility that 
queens of this species can produce a work- 
er from a single foraging trip. 

The male H. lanei is easily distinguished 
from H. lutescens and H. hesperus, and in 
fact all other Seladonia, by the unique gen- 
italic and sternal characters, though again, 



Figs. 34-37. Halictufi terminalia. 34-36. H. hvici. 34, Sterna 7 and 8. 35, Genital capsule, dorsal view (left), 
ventral (right). Dorsal and ventral views shown at angles optimizing view of relevant structures, not sym- 
metrical. 36. Gonostylus, lateral view. 37. H. (S.) Iicspcrut^, gonostylus, lateral view. Hairs and portions of hairs 
behind other structures not drawn. 



74 



Journal of Hymenoptera Research 



only H. hesperus has a geographic range 
approaching that of H. lanei. In addition to 
the unique features of H. lanei, males of 
the two species can be distinguished by 
the following characters: H. hesperus has 
the anterior punctures on the scutum 
stronger and more effaced than H. lanei, 
with the scutum also more swollen ante- 
riorly on either side of the median suture 
such that it appears biconvex and the 
punctures of the scutum and scutellum 
are more dense in H. hesperus than H. lanei. 
Halictus hesperus has darker legs, especial- 
ly the coxae and trochanters, which are all 
brown in this species. S6 of H. Jiesperus has 
a shallow, ill-defined, impressed, glabrous 
area which does not reach the apex of the 
sternum and is separated from the apex 
by a region which is hirsute like the lateral 
areas. Conversely, in H. lanei the impres- 
sion on S6 is deep, with a well defined, 
transverse anterior margin, and it reaches 
the apex of the sternum (Fig. 33). The gen- 
italia also differ between the two species. 
In dorsal view, the gonobase of H. lanei is 
% as long as it is wide, whereas in hespe- 
rus, it is less than X; as long as wide, and 
is much more rounded; the gonostylus of 
H. hesperus is more quadrate, with a less 
pronounced dorso-basal projection and 
has a stronger swelling ventrally; this spe- 
cies also has denser hair on the dorsally 
recurved part of the gonostylus (compare 
Figs. 36 and 37); and the medially directed 
apico-dorsal projection of the penis valve 
is more pronounced in H. hesperus than in 
H. lanei. 

The hair patch on the fifth sternum of 
the male is reminiscent of a similar feature 
in many species of the subgenus Vestito- 
halictus. However, in Vestitohalictus the 
hair tuft is most commonly found on the 
fourth sternum (Michener, 1978), although 
in some species, such as H. (V.) concinnus 
it is repeated on the fifth sternum (Packer, 
unpublished data). Owing to the pre- 
sumed phylogenetic position of these sub- 
genera and species, it is unlikely that this 



hair tuft is homologous between the two 
subgenera. 

ACKNOWLEDGMENTS 

We would like to thank the following for helping 
us obtain specimens required for this study or to ob- 
tain further information about the specimens: the late 
Professor George Eickwort of Cornell University; Dr. 
Terry Griswold of the USDA bee lab, Logan, Utah; 
David Hawks and Dr. Douglas Yanega of the Uni- 
versity of California, Riverside; Dr. Albert Finnamore 
of the Provincial Museum of Alberta; Dr. Jon Gelhaus 
of the Philadelphia Academy of Sciences; Dr. John 
Rawlins of tlie Carnegie Museum and Dr. Danuncia 
Urban of Curitiba. Dr. Bryan Danforth kindly sup- 
plied drawings of S7 and S8 of a male of H. har)iioiuus 
before we found additional specimens in the collec- 
tion at Riverside. We thank Drs. Mike Engel and Eric 
Grissell for helpful comments on the manuscript. Our 
research was fundeci by an NSERC graduate schol- 
arship to the senior author and an NSERC research 
grant to the junior author. 

LITERATURE CITED 

Brooks, R. W. and D. W. Roubik. 1983. A halictine 
bee with distinct castes: Halictus hesperus (Hy- 
menoptera: Halictidae) and its bionomics in cen- 
tral Panama. Sociobiology 7:263-282. 

Eickwort, G. C. 1969. A comparative morphological 
study and generic revision of the augochlorine 
bees (Hymenoptera: Halictidae). University of 
Knnsns Science Bulletin. 48:325-524. 

Krombein, K. V., P. D. Hurd Jr., D. R. Smith, and B. 
D. Burks. 1979. Catalog of the Hyiiieiioptera of 
Aiuerica North of Mexico, vol. 2, Apocrita. Smith- 
sonian Institution Press, Washington, xvi -I- pp. 
1199-2209. 

McGinley, R. J. 1986. Studies of Halictinae (Apoidea: 
Halictidae), I. Revision of New World Lasioglos- 
suin Curtis. Siiiithsoiiiaii Contributio)is to Zoology, 
No. 429. 294pp. 

Michener, C. D. 1954. Bees of Panama. Bulletin of the 
American Museum of Natural History. 104:1-175. 

Michener, C. D. 1978. The classification of halictine 
bees: tribes and Old World non-parasitic genera 
with strong venation. University of Kansas Seieiwe 
Bulletin 51:501-538. 

Michener, C. D., R. J. McGinley, and B. N. Danforth. 
1997. The bee genera of North ami Central America. 
Smithsonian Institution Press, Washington, 
viii + 209pp. 

Moure, J. S. 1940. Apoidea neotropica. Arquivos do 
Zoologia Sao Paulo. 2:36-64. 

Packer, L. 1985. The social organisation of two hal- 
ictine bees from southern Mexico with notes on 
two bee-hunting philanthine wasps. Pan-Pacific 
Entomologist 51:291-298. 



Volume 10, Number 1, 2001 



75 



Packer, L., and J. S. Taylor. 1997. How many cryptic 
species are there? An application of the phylo- 
genetic species concept to genetic data for some 
comparatively well known bee species. Caiiadimi 
Entomologist 129:387-594. 

Rosenmeier, L., and L. Packer. 1993. A comparison of 
genetic variation in two sibling species pairs of 
haplodiploid insects. Biochemical Genetics 31:185- 
200. 

Sandhouse, G. A. 1941. The American species of the 



subgenus Halictus. Entomologicn Americnna, New 
Series 21:23-38. 

Walker, K. 1995. Revision of the Australian native bee 
subgenus Lasioglossiim (Chitnlictus) (Hymenop- 
tera: Halictidae). Memoirs of the Museum of Vic- 
toria 55:1-423. 

Wille, A., and C. D. Michener. 1971. Observations on 
the nests of Costa Rican Halictus with taxonomic 
notes on Neotropical species (Hymenoptera: Hal- 
ictidae). Rei'ista de Biologia Tropicnle 18:17-31. 



J. HYM. RES. 
Vol. 10(]), 2001, pp. 76-80 

The New Western Australian Tiphiid Genus Dythynmis Kimsey 
(Hymenoptera: Tiphiidae: Thynninae) 

Lynn S. Kimsey 

Bohart Museum of Entomology, Department of Entomology, University of California, 

Davis, CA 95616 



Abstract. — The new genus Dythynnus Kimsey is described from Western Australia based on 
the new species Solaris Kimsey and thermophilus Kimsey (type of genus). 



Northern and central Western Australia 
are among the most poorly collected re- 
gions in Australia. A large number of 
thynnine species from this area fail to fit 
into established genera. Although the pre- 
cise relationships betu^een Dythynnus 
Kimsey and other Australian thynnine 
genera remain unresolved, preliminary 
phylogenetic analyses of these genera sug- 
gest that Dythynnus is at least a basal lin- 
eage of a clade containing genera related 
to Iswaroides Ashmead, as discussed by 
Kimsey (1999). This is based upon a num- 
ber of male characteristics including the 
laterally notched transverse pronotal cari- 
na (welt), the epipygium subapically with 
a partial transverse carina, volsella U- 
shaped in cross-section, gonobase narrow- 
ly attached to gonocoxa, and slender penis 
valves. This genus also has the palpal 
brush seen in Chilothynnus Brown. 

Specimens were obtained through ex- 
tensive collecting in Western Australia. 
Both holotypes will be placed in the West- 
ern Australian Museum, Perth. Paratypes 
will be dispersed between the Australian 
National Insect Collection, CSIRO, Can- 
berra, ACT, the Bohart Museum of Ento- 
mology, University of California, Davis, 
USA, and the Western Australian Muse- 
um. The terms hypostomal plate and hy- 
postomal carina are used in the sense of 
Bohart and Menkc (1976). 



Dythynnus Kimsey, new genus 

Male. — Body length 6-9 mm. Head: clyp- 
eus narrowly truncate apically, truncation 
round cornered and 1-2 midocellus di- 
ameters across; antennal lobes small, 
closely aligned and rounded, without 
ridge or carina, strongly elevated above 
subantennal sclerite; frons with frontal 
line extending nearly to midocellus; la- 
brum small and apically bilobate; vertex 
without red spot behind hindocellus; basal 
maxillary palpal segment with long, erect 
setae; hypostomal plate extending to outer 
mandibular socket; occipital and hypos- 
tomal carinae narrowly to broadly sepa- 
rated ventromedially; prementum asetose, 
longitudinally grooved laterally; stipes 
with sparse marginal fringe, traversing sti- 
pes at midpoint; flagellomere I 1.5-2X as 
long as broad; flagellomere II 2.5-3X as 
long as broad; flagellomeres without ty- 
loids or with single, often indistinct, sub- 
apical one. Mesosoma: pronotal disk ante- 
rior margin marked by transverse swell- 
ing or broad ridge, without medial or sub- 
lateral indentations or notches; scrobal 
sulcus shallow, extending less than half 
way across mesopleuron; propodeum 
sloping obliquely from metanotum to pet- 
iolar socket; forecoxa globular to flat, me- 
dially setose to asetose; legs unmodified 
except hindtrochanter produced posteri- 
orly into sharp apical angle or tooth. Mc- 
tasoma: tergum I about as broad as long. 



Volume 10, Number 1, 2001 



71 



gently convex subapically, sternum I me- 
dially convex; terga I-VI and sterna II-V 
each with subapical transverse sulcus; ter- 
ga I-V each with subspiracular sulcus; epi- 
pygium with thin subapical transverse ca- 
rina or none, with thin to broad apical 
transparent rim; hypopygium narrowly 
triangular with transparent lateral edge. 
Genital capsule (as in Figs. 3-5, 8-10): gon- 
ocoxa dorsoapically lobate; gonobase nar- 
rowly attached to gonocoxa in lateral 
view; paramere apically bilobate or trun- 
cate; aedeagus with long apical loop; vol- 
sella large, U-shaped in cross section, with 
slender inner and outer lobes (as in Figs. 
6, 11); penis valves slender apically. Color: 
black, strongly marked with yellow, or- 
ange and red. 

Female. — Unknown. 

Typ^e species. — Dythynnus theniiophilus 
Kimsey, new species. 

Etymology. — The generic name is a non- 
sense combination of letters added to the 
commonly used suffix in this tribe — 
"thynnus". The name is assumed to be 
masculine. 

Included species. — Dythynnus Solaris Kim- 
sey, new species and Dythynnus tJiermo- 
philus Kimsey, new species. 

Distribution. — This genus is known from 
two regions in Western Australia, the Stir- 
ling Range (Solaris) and 100-1- km north of 
Northampton {thermophilus), in the sum- 
mer months of December and January. 

Discussion. — Dythynnus species can be 
immediately separated from other Austra- 
lian genera by the sparse, arcuate stipal 
fringe, occipital and hypostomal carinae 
broadly separated, flagellomeres without 
or with only a single tyloid, laterally 
notched pronotal welt, epipygium with 
broad, polished and impunctate apicome- 
dial area, and basal maxillary palpal seg- 
ments with erect brush-like setae. The 
brushy palpal segments are similar to 
those seen in Chilothynnus Brown. How- 
ever, the closely aligned antennal sockets 
(about 1 midocellus diameter apart), elon- 
gate and narrowly triangular hypopy- 



gium, and arcuate stipal fringe will im- 
mediately separate Dythynnus from Chilo- 
thynnus. 

Dythynnus Solaris Kimsey, new species 

(Figs. 1, 3-7, 12) 

Male. — Body length 6-7 mm. Head (Fig. 
1): facial punctation contiguous to 1 punc- 
ture diameter apart, punctures becoming 
larger on frons; vertex with punctures be- 
hind ocelli 1-2 puncture diameters apart; 
clypeal apex 0.2 X greatest width of clyp- 
eus; clypeus greatest width 2.6 X length; 
face 1.2X as long as greatest width above 
antennal sockets; midocellus 4.2 midocel- 
lus diameters from nearest eye margin; 
flagellomere I 1.7X as long as broad; fla- 
gellomere II twice as long as broad; fla- 
gellomere III 2.5 X as long as broad. Me- 
sosoma: pronotal and scutal punctures 1-2 
puncture diameters apart; scutellar punc- 
tures 2-4 puncture diameters apart; me- 
tanotum nearly impunctate; propodeal 
punctures nearly obscured by dense, fine, 
transverse ridges, smooth and impunctate 
anteriorly; mesopleural punctures 0.5-1 
puncture diameter apart; propleuron con- 
vex and setose; forecoxa convex, covered 
with long setae, setae becoming densest in 
medial patch; Metasoma: terga and sterna 
highly polished, punctures small and shal- 
low, 2-4 puncture diameters apart; epi- 
pygium subapically narrowed with broad 
transparent rim; hypopygium (Fig. 12). 
Genital capsule (Figs. 3-5): paramere broad 
and apically truncate in lateral view, apex 
bending toward midline, dorsally densely 
setose; gonocoxa dorsally broadly bilo- 
bate, dorsal surface depressed sublateral- 
ly; volsella with slender digitate inner lobe 
and broader, acute outer lobe (Fig. 6). Col- 
or: head black, with whitish band along 
eye margin and becoming reddish behind 
vertex; clypeus and mandible whitish; 
frons with short whitish band above each 
antennal socket; occiput and hypostoma 
black; pronotum black, with transverse 
anterior and posterior whitish bands; scu- 
tum and scutellum black, with whitish 



78 



JOURNAI. OF HYMENOPTERA RESEARCH 




thermophilus thermophilus thermophilus 



13. thermophilus 



Figs. 1-13. D\/tlnpiini> species. 1, 2, FrDiit view of male Kice, left antenna not shown. 3, 8, Dorsal view of 
genital capsule. 4, 9, Lateral view of genital capsule. 5, 10, Ventral view of genital capsule, with gonobase 
removed. 6, 11, Ventral view of volsella. 7, Ventral view of male head, with occipital region not shown. 12, 
13, Dorsal view of metasomal segment VII, tergum and sternum. 



Volume 10, Number 1, 2001 



79 



medial and lateral spots; metanotum 
black, with whitish medial spot; propo- 
deum black; mesopleuron black, with 
whitish comma-shaped anterior and 
rounded posterior spots; propleuron 
black; forecoxa black, with large medial 
whitish spot and interior black spot (cor- 
responding with tuft of long setae); fore- 
legs red; mid and hindlegs, coxae black 
becoming whitish dorsally, trochanters 
brown, femora red, tibiae red ventrally, 
brown dorsally, tarsi brown; metasomal 
tergum II black, with pale lateral spot; ter- 
ga III-Vl reddish brown, becoming darker 
posteriorly, with pale whitish lateral spot; 
terga VII-VIII dark brown to black; ster- 
num II black; sterna III-VII red; sternum 
VIII dark brown; wing veins brown, be- 
coming pale basally; wing membrane un- 
tinted. 

Type viaierial. — Holotype 6: 7 km n Stir- 
ling Range, 34°19'S 118°11"E, 23-25 Dec. 
1994, L. S. & R. B. Kimsey, ex Eucalyptus 
flowers, field No. WA122401 (PERTH). 
Paratypes: 1 6 : same data as holotype; 1 
6 : 24 Dec. 1994; 1 6 : 23-26 Dec. 1994; 3 
6 : 24-26 Dec. 1994, field No. WA122305 
(CANBERRA, DAVIS, PERTH). 

Etymology.— The species name means 
"of the sun", Latin, masculine; it refers to 
the intense summer conditions when the 
males fly. 

Discussion. — Dythynnus Solaris can be 
distinguished from thermophilus by the 
apically truncate parameres, flagellomere 
I less than twice as long as broad, pale, 
whitish markings, and entirely black pro- 
podeum. 

Dythynnus themiophihis Kimsey, new 
species 

(Figs. 2, 8-11, 13) 

Male. — Body length 6-9 mm. Head (Fig. 
2): face highly polished; subantennal scler- 
ite impunctate; clypeal punctures 3-5 
puncture diameters apart; area between 
antennal socket and eye nearly impunc- 
tate; frons and vertex highly polished, 
punctures 2-8 puncture diameters apart; 



clypeal apex 0.2 X greatest width of clyp- 
eus; clypeus greatest width 2.6 X length; 
face 1.3-1.4X as long as greatest width 
above antennal sockets; midocellus 2.8 mi- 
docellus diameters from nearest eye mar- 
gin; flagellomere I 2.2 X as long as broad; 
flagellomere II 3 X as long as broad; fla- 
gellomere III 3.2-3.5 X as long as broad. 
Mesosomn: pronotum, scutum and scutel- 
lum highly polished, punctures 4-8 punc- 
ture diameters apart, becoming denser lat- 
erally; metanotum nearly impunctate; pro- 
podeum impunctate medially, punctures 
1-2 puncture diameters apart laterally; 
mesopleuron highly polished, punctures 
2-4 puncture diameters apart; propleuron 
convex and evenly setose; forecoxa slight- 
ly flattened and nearly asetose in most 
specimens. Metasoma: tergal punctures 
tiny and obscure, 1^ puncture diameters 
apart anteriorly, markedly larger and 
denser on tergum VII; sternal punctures 
1-4 puncture diameters apart; epipygium 
with thin, transverse, subapical carina, 
and narrow, transparent apical rim; hy- 
popygium (Fig. 13). Genital capsule (Figs. 
8-10): paramere strongly bilobate, with 
acute dorsal lobe and elongate digitate 
ventral one, nearly asetose; gonocoxal dor- 
sal lobes broadly rounded, overall appear- 
ing shallowly bilobate dorsomedially; vol- 
sella with long digitate outer lobe and 
much shorter, acute inner lobe (Fig. 11). 
Color: Head yellow, except frons midline 
brown to black in some specimens, and 
black line extending dorsally above anten- 
nal sockets connecting with line forming 
box around ocelli; some specimens with 
brown line extending from posterior eye 
margin and across vertex behind ocelli; 
occiput black; pronotum yellow except 
transverse medial stripe and anterior face; 
scutum black, with large tridentate medial 
spot and yellow laterally; scutellum yel- 
low, with black anterior margin connected 
to sublateral black band; metanotum yel- 
low medially and anteriorly, black later- 
ally; propodeum black, with broadly tri- 
angular yellow medial spot and large 



80 



Journal of Hymenoptera Research 



comma-shaped lateral yellow spot; pro- 
pleuron black, with large yellow medial 
spot; coxae yellow, with black base; mid 
and forelegs yellow and red, with apical 
tarsomeres brown (tibiae and femora may 
be entirely yellow, and foretarsi yellow in 
some specimens); hindfemur red, yellow 
laterally, hindtibia red, hindtarsomeres 
brown; mesopleuron black, with most or 
all of lateral surface yellow, scrobal sulcus 
marked by black and brown in some spec- 
imens; metasomal tergum II black basally, 
yellow submedially and red posteriorly; 
terga III-VI each red becoming brown 
posteriorly, each with large yellow lateral 
spot or complete yellow band; metasomal 
sterna red, except sternum VIII dark red 
to black; wing veins pale brown, becom- 
ing paler basally; wing membrane untint- 
ed. 

Ti/pe ninterial. — Holotype 6: 115 km n 
Northampton, 9 Jan. 1995, 27°27'S 
114°41'E, R. B. & L. S. Kimsey, ex Eucalyp- 



tus flowers, field No. WA010905 (PERTH). 
Paratypes: 19 6: same data as holotype; 9 
6: 120 km n Northampton, 27°25'S 
114°40'E, R. B. & L. S. Kimsey, ex Eucalyp- 
tus flowers, field No. WA010904 (CAN- 
BERRA, DAVIS, PERTH). 

Etymology. — The name, "therme", "phi- 
lus" , means heat-loving, Greek, masculine; 
it refers to the hot summer conditions en- 
countered when the specimens were col- 
lected. 

Discussion. — This species can be readily 
distinguished from Solaris by the strikingly 
bilobate parameres, much longer first fla- 
gellomere and bright yellow markings, 
particularly the yellow medial propodeal 
spot. 

LITERATURE CITED 

Bohart, R. M. and A. S. Menke. 1976. Sphccid ivasps of 
the world. University of California Press, Berke- 
ley, ix + 695 pp. 

Kimsey, L. S. 1999. What is the real Iswaroidcs? Pro- 
ceedings of the Wiishiiigtoji E)itoiiiologicid society 
101:503-513. 



J. HYM. RES. 
Vol. 10(1), 2001, pp. 81-90 

Effects of Parasitism by Banchus flavescens 

(Hymenoptera: Ichneumonidae) and Microplitis mediator 

(Hymenoptera: Braconidae) on the Bertha Armyworm, Mamestra 

configurata (Lepidoptera: Noctuidae)' 

P. G. Mason-, M. A. Erlandson, and B. J. Youngs 

Agriculture and Agri-Food Canada, Saskatoon Research Centre, 107 Science Place, 
Saskatoon, Saskatchewan S7N 0X2 



Abstract. — The bertha armyworm, Mamestra configurata Walker (Lepidoptera: Noctuidae), is an 
important pest of canola {Brassica iiapms L. and B. rapa L.) and flax {Limini usitatissimum L.) in 
western Canada. It is the focus of research to develop a management program integrating micro- 
bial and insect parasitoid agents with cultural practices and judicious use of chemical insecticides. 
To develop IPM effectively it is essential to understand the impact of parasitism on feeding activity 
of the host. We studied the effect of parasitism by two solitary koinobiont larval endoparasitoids: 
Ba7ichus flavescois Cresson (Ichneumonidae) is a native parasitoid of M. configurata; and the Eu- 
ropean Microplitis mediator (Haliday) (Braconidae) is a candidate for introduchon to enhance the 
biological control effected by B. flavescens. Parasitism by B. flavescens resulted in significantly de- 
creased food consumption and lower biomass production but did not reduce the time that the 
pest would occur in the crop. Host larvae parasitized by M. mediator showed a much greater 
reduction in food consumed, weight gained, trass produced and the host's feeding time compared 
to nonparasitized larvae or those parasitized by B. flavescens. Management strategies should con- 
sider options that would minimize impact on parasitized larvae and introduction of M. iiwdiator 
could benefit integrated management programs for M. configurata. 



The bertha armyworm, Mmiwstra coiifi- 
giirntn Walker, native to North America, has 
been the focus of studies to determine the 
role of natural control agents for this pest of 
canola {Brassica napus L. and B. rapa L.: Bras- 
sicaceae) and flax (Linuni usitatissimum L.: 
Linaceae), the two most important oilseed 
crops grown in western Canada. Important 
natural control agents of larval stages of the 
bertha armyworm include viral and fungal 
diseases and parasitic wasps and flies (Wy- 
lie 1977, Wylie and Bucher 1977, Arthur and 
Mason 1985, Turnock 1988, Erlandson 
1990). The solitary koinobiont ichneumonid 
Banchus flavescens Cresson is the single most 



' Saskatoon Research Centre Contribution No. 1272. 
' Present address. Agriculture and Agri-food Canada, 
Eastern Cereal and Oilseed Research Centre, Ottawa, 
Ontario, KIA 0C6 Canada. 



important larval parasitoid, occurring in 
more than 90% of bertha armyworm sam- 
pled in years when populations are declin- 
ing (Turnock and Bilodeau 1984, Arthur 
and Mason 1985) yet B. flavescens is unable 
to prevent outbreaks from occurring (Ma- 
son et al. 1998). 

Biologically based pest management 
strategies being developed for the bertha ar- 
myworm include microbial insecticides (nu- 
clear polyhedrosis virus, MacoNPV (Erland- 
son 1990)) and introduction of exotic para- 
sitoids to complement the mortality caused 
by native natural enemies. It is essential to 
understand the impact of parasitoids on the 
host's feeding activity so that management 
strategies can be developed to minimize the 
impact on parasitoids. 

The solitary koinobiont braconid parasit- 
oid Microplitis mediator (Haliday) has been 



82 



Journal of Hymenoptera Research 



Studied as a candidate for introduction to 
enhance natural biological control of M. con- 
figumta. M. medintor attacks the cabbage 
moth, Mamestrn brassicne (L.), in its native 
habitat in Europe (Slovak 1985a, 1985b, Jo- 
hansen 1997). It is synchronized with its 
host in both uni- and bivoltine systems. Ar- 
thur and Mason (1986) demonstrated that 
M. mediator will attack and successfully par- 
asitize 1"' to 3'^' instar bertha armyworm lar- 
vae, although 3'^' instars frequently repel or 
injure female wasps attempting to oviposit. 
The fully developed parasitoid larva egress- 
es from the host's 4"' instar, regardless of the 
instar parasitized (Mason, unpublished) 
while non-parasitized larvae complete 6 in- 
stars and then pupate. 

Evaluation of the impact of candidate 
biological control agents on the target eco- 
system is a necessary component of bio- 
logical control screening projects. An im- 
portant part of this process is to determine 
the effect of the agent on the crop being 
protected. Although idiobiont parasitoids 
normally kill the host immediately (elim- 
inating further plant feeding) koinobiont 
insect endoparasitoids generally do not 
kill immediately and permit feeding by 
the host for at least a short period of time 
after parasitization (Askew and Shaw 
1986, Quicke 1997). Thus, determination of 
the effect of koinobiont parasitisn^i on the 
host's feeding is an indirect means for de- 
termining the impact of parasitoids on the 
crop being protected. Depending on par- 
asitoid biology, parasitism can reduce 
(Rahman 1970, Guillot and Vinson 1973, 
Brewer and King 1978, Sajap et al. 1978, 
Brewer and King 1980, Parkman and 
Shepard 1981, Shaw 1981, Powell 1989, 
Kumar and Ballal 1992, Ohnuma and Kai- 
noh 1992, Khan 1994, Yang et al. 1994, 
Harvey 1996), increase (Rahman 1970, 
Hunter and Stoner 1975, Byers et al. 1993) 
or have no effect on (Brewer and King 
1981) feeding by lepidopteran host spe- 
cies. Parasitoids that elicit decreased feed- 
ing by the host reduce potential damage 
to the crop in addition to removing the 



host from the reproductive pool for sub- 
sequent generations. 

The application of insecticides for con- 
trol of bertha armyworm usually takes 
place during the pod development stage 
of canola, when the larvae are in the 5* 
and 6"' instars. The decision to apply in- 
secticides is based on evidence of real or 
potential (often visibility of larvae) crop 
damage. Because insecticides may have a 
negative impact on adult and larval par- 
asitoids, understanding parasitoid biology 
is important to develop sound pest man- 
agement strategies. Arthur and Mason 
(1985, 1986) showed that B. flavescens and 
M. mediator oviposit in 1"' to 3'^'^* instar ber- 
tha armyworm which are inconspicuous 
and feed on canola leaves before pods de- 
velop. Because no control measures are 
taken during these life stages of the pest, 
adult B. flavescens and M. mediator are not 
exposed to insecticides when they are 
searching in the foliage of the canola crop. 
Further, they are unlikely to be present 
when insecticides are applied during the 
5"' and 6"' larval instars because the pre- 
ferred instars for oviposition are not pres- 
ent. In contrast, parasitized host larvae 
may still be present when insecticides are 
applied. For example, parasitoid species 
that prolong development of the host by 
increasing or making no difference in the 
amount of food consumed by the pest are 
more likely to be indirectly exposed to and 
negatively impacted by control measures 
such as chemical insecticides that kill on 
contact and are usually used against ma- 
ture larvae. Parasitoid species that reduce 
food consumption are less likely to be af- 
fected because they tend to shorten the life 
of the host and reach a life stage (i.e. form 
protective cocoons) in which insecticides 
may not kill them. Thus, the impact of 
parasitism on the host's feeding may affect 
plans for biological control introductions 
in integrated pest management (IPM) pro- 
grams (Hunter and Stoner 1975). In this 
study we assessed the effect of parasitism 
by the native ichneumonid B. flavescens 



Volume 10, Number 1, 2001 



83 



and the exotic braconid M. medintor on M. 
configurata growth and food consumption. 

MATERIALS AND METHODS 

Hosts used in the experiments were 
from a M. coufiguratn colony maintained 
on artificial diet (Bucher and Bracken, 
1976). Parasitoids used in the experiments 
were derived from populations collected 
in Saskatchewan, Canada (B. flavescens) 
and central Switzerland (M. mediator). The 
B. flavescens females used in the experi- 
ments were reared from overwintered co- 
coons collected in the spring and main- 
tained at 21±1°C and 16:8 L:D. The M. 
mediator was maintained in laboratory cul- 
ture on M. configurata (Arthur and Mason, 
1986). All plants used were canola, Bras- 
sica napns L. variety Westar. All experi- 
mental individuals were maintained at 
21±1°C and 16:8 L:D. 

For each of the experimental trials (3 for 
B. flavescens and 4 for M. mediator), 30 2nd 
instar larvae from the same egg mass were 
selected. The larvae were divided into 
three groups of 10 and individuals of each 
group weighed. Whole canola leaves (ap- 
proximately 7x6 cm) clipped at the pet- 
iole from growth cabinet grown plants 
were used in all food offerings and dry 
weight determinations. Larvae of one 
group were placed in individual 52mm di- 
ameter X 21mm deep petri dishes contain- 
ing a canola leaf on moistened filter paper 
in 45 mm diameter petri dishes. Larvae of 
a second group were individually parasit- 
ized (under observation) by 6. flavescens or 
M. mediator and placed on canola leaves in 
the manner described above. The third 
group of larvae were individually 
weighed (wet weight) with a Mettler an- 
alytical balance (Model AG245), frozen, 
dried (48 h @ 60°C), and then individually 
weighed (dry weight). The larvae were 
weighed and given fresh leaves at inter- 
vals (2-4 day) during each trial. The fresh 
weight of each leaf disc was recorded be- 
fore introduction into the petri dish. Fresh 
and dry weights (48 h @ 60°C) were re- 



corded separately and dry weight ratios 
(dry weight ^ initial fresh weight) were 
calculated. The frass produced by each 
larva was collected at the end of each 
feeding period and stored in a plastic petri 
dish. The pooled amount for the experi- 
mental period was dried (48 h @ 60°C) and 
weighed. Similarly the fresh:dry weight 
ratios were determined for all larvae after 
parasitoid egression for parasitized indi- 
viduals and at pupation for non-parasit- 
ized larvae. 

Nutritional indices were initially con- 
sidered (Waldbauer 1968, Kogan 1986), 
but Schmidt and Reese (1986) pointed out 
that these indices can be misleading due 
to the error introduced during initial mea- 
surements of dry matter of the food 
source, quantity of food eaten, and the 
weight of feces produced. These errors ac- 
cumulate and are amplified in the calcu- 
lation of the nutritional indices (Rauben- 
heimer and Simpson 1992). Van Loon 
(1991) describes more precise techniques 
including indigestible markers, elemental 
budgets, planimetry, and respirometry. 
Thus, we calculated only: 

Food consumption (F) = [(Mean Dry Weight 
of Control Food Portions /Mean Wet 
Weight of Control Food Portions) X Ini- 
tial Wet Weight of Food Portions] - 
Dry Weight of Food Portions after feed- 
ing; 

Total Food Consumed (TC) = iS"F, where n 
is the number of feeding intervals (to 
parasitoid egression or host pupation); 

Total Dry Weight Produced (DWT) = Dry 
Weight at host pupation or parasitoid 
egression (host + parasitoid) - Mean 
Dry Weight (of control group) at the be- 
ginning of the experiment. 

The B. flavescens data were analysed us- 
ing analysis of variance (Proc GLM, SAS 
1992) to determine if parasitism signifi- 
cantly affected the variables measured. 
The effects of parasitism by M. mediator 
were so obvious that statistical analyses 



84 



Journal of Hymenoptera Research 



Table 1. Mean value for development time (days){DT), total dry w^eight (mg)(ot pupa or parasite larva + 
larval skin produced )(DWT), total mg dry weight of trass produced (Frass), and total mg dry weight consumed 
(TC) for non-parasitized and B. flnvcscois-parasitized M. avifi^iirata. 





Trie 


d#i 




Ti 


-lal #2 






Irial #3 






[^ooled 


Indfx 


\P 




r 


\P 




P 


\p 




P 


Mean 


SEM 


DT 


21.4* 




21 


19.2 




19.3 


20.1 




21 


20.3 


0.2 


DWT 


83.2 




32.8' 


57.2 




32.3^ 


80.2 




46.7^ 


53.3 


3.5 


Frass 


142.9 




117.3 


185 




127.0'^ 


138 




166.9 


148.2 


6.1 


TC 


415.3 




318.9' 


406.5 




228.4' 


371 




297.6^ 


334.9 


12.6 



^approximate standard error of the mean for a trial mean can be obtained by multiplying the pooled SEM by 1.7 
For each trial significant differences owing to parasitism are indicated by: ' = (P < 0.001), '-^ = {P < 0.01), 
' = (P < 0.1). 



were not necessary (a non-parametric test 
could have been used). 

RESULTS 

Banchus flavescens. — In all experimental 
trials, nonparasitized (NP) M. configurata 
larvae consumed significantly more food 
and gained significantly more weight than 
did parasitized (P) larvae (Table 1). There 
was no significant difference in the num- 
ber of days required to complete devel- 
opment. From the beginning of the trial it 
took M. configurata 18-24 days to develop 
to the pupal stage and development of B. 
flavescens from oviposition to larval egres- 
sion was completed in 19-24 days. How- 
ever, in spite of minor differences among 
trials on some dates, overall the only var- 
iable which showed a significant trial X 
treatment interaction, that is differences in 
pattern from trial to trial, was the dry 
weight of frass produced. 



While Fig. la and b are representative 
of the patterns observed in all three trials, 
owing to individual variability within tri- 
als (especially because some values were 
much higher or lower than all others in a 
group) differences in weight and food 
consumed by the parasitized and non-par- 
asitized groups were not always consis- 
tent, on specific dates, from trial to trial. 
Consumption of food peaked at 14-18, 16- 
19 and 11-14 days in trials 1-3, respec- 
tively. Food consumption declined mark- 
edly after the peak, coinciding with the 
beginning of the prepupal stage in non- 
parasitized hosts and the pre-emergence 
period of the parasitoid. 

Micwplitis mediator. — In all experimental 
trials non-parasitized (NP) M. configurata 
larvae consumed dramatically more food, 
gained niuch more weight and produced 
much more frass than did parasitized (P) 
larvae (Table 2). Feeding and develop- 



Tahle 2. Mean value for development time (DT), total dry weight (of pupa or parasite larva + larval skin 
produced (DWT)), total mg dry weight of frass produced (Frass), and total mg dry weight consumed (TC) 
for non-parasitized and M. »/('(/;(?f(ir-parasitized M. coiifi^uiata. 





In, 


il #: 




T: 


rial #. 


2 


Ir 


lal #3 


1 


rial #4 


Pool. 


.•d SE 


Indev 


\l' 




p 


\P 




P 


\P 


P 


\i' 


P 


\p 


1' 


DT 


21 




12 


21 




11.4 


21 


12.3 


21 


11 





0.2 


DWT 


67.6 




2.8 


62.4 




1 


77.8 


3.8 


81.2 


2.6 


4.2 


0.02 


Frass 


182.6 




5.1 


209 




4.5 


161 


7 


213 


12.7 


8.3 


0.1 


TC 


402 




13 


593 




82 


294 


31 


356 


47.6 


26.1 


13.9 



approximate standard error of the mean for a trial mean can be obtained by multiplying the pooled S.E.M. 
2.0. 



by 



Volume 10, Number 1, 2001 



85 



ment to the pupal stage of non-parasitized 
M. configiirata reared from neonate larvae 
at 21±rC takes 21-25 days (unpublished 
data). Parasitized larvae ceased to con- 
sume food, once M. riiedintor larvae 
egressed and formed cocoons (11-14 days 
after parasitism), a markedly shorter pe- 
riod than the food consumption period for 
non-parasitized M. coufigiirnta larvae. 

Non-parasitized larvae weighed and 
consumed significantly more beginning 4- 
8 and 8-11 days, respectively, after trials 
were initiated. The trends represented in 
Figure 2a and b for trial #1 were typical. 
For non-parasitized larvae consumption of 
food and larval weights peaked 15-17, 13- 
15, 11-13, and 13-15 days after parasitism 
in trials 1-4, respectively. Both consump- 
tion and weight declined after this peak, 
coinciding with the beginning of the pre- 
pupal stage. For parasitized larvae, larval 
weight did not increase beginning 4-8 
days after the trials were initiated. Food 
consumption showed only a small in- 
crease in trial #2, while actually decreasing 
in the other three trials beginning 0-4 days 
after trials were initiated. In all trials food 
consumption by parasitized larvae ceased 
by day 13, 7 days before non-parasitized 
larvae ceased feeding. 

DISCUSSION 

Synchronization of the completion of 
larval development of B. flavescens and M. 
configuratn is consistent with results by 
Slovak (1987) for another Banchini, Exes- 
tastes cinctipes Retzius (Hymenoptera: Ich- 
neumonidae), parasitizing M. brassicae. He 
found that at 20°C development to para- 
sitoid cocoon formation took 25.3-26.6 
days while non-parasitized larvae took 
26.4—26.7 days to form pupae. These de- 
velopment times are 5-6 days longer than 
those we observed at 21.0 ± 1.0°C for B. 
flavescens parasitizing M. cotifigurata. Slo- 
vak (1987) also noted that at 20 or 24°C 
parasitized host larvae entered the soil 
significantly earlier (2.1-3.0 and 3.5-3.9 
days, respectively) than non-parasitized 



larvae. Although we did not study this be- 
haviour, it may also occur in the B. flaves- 
cens /M. configiirata parasitoid/host sys- 
tem because, as reported by Arthur and 
Mason (1985), the parasitoid forms a co- 
coon in which it overwinters within the 
earthen cell formed by the host larva just 
before being killed. 

That parasitism by B. flavescens signifi- 
cantly reduced the total amount of food 
consumed by M. configiirata (20^4%) is 
reflected in the fact that parasitized larvae 
produced significantly less (39.4-58.2%) 
dry protein matter than non-parasitized 
larvae. Although not directly comparable, 
these results are siniilar to findings for sol- 
itary koinobiont Campoplegine species at- 
tacking Lepidoptera (Khan 1994, Kumar 
and Ballal 1992, Yang et al. 1994, Doucet 
and Cusson 1996). Only Doucet and Cus- 
son (1996) measured frass production, and 
found that non-parasitized larvae pro- 
duced 38% more frass than parasitized lar- 
vae. In contrast, about 10% more frass was 
produced by non-parasitized than parasit- 
ized larvae in two of three trials and 10% 
less in the third trial in our study. It is not 
clear why frass production by parasitized 
larvae in one of our three trials was great- 
er than for nonparasitized larvae. 

Development of M. mediator to egres- 
sion took 11-14 days, about 1/3 less time 
than for B. flavescens. Egression of mature 
M. mediator larvae occurs just after the 
host molts to the 4"' larval instar even if 
parasitized in the 3"^^^ instar and, as noted 
by Shaw and Huddelston (1991) for other 
Microplitis spp., the cocoon is formed be- 
neath the host which remains alive for 
several days without feeding (Mason, un- 
published). Behavioral modification of M. 
cotifigurata hosts by mature larval M. me- 
diator parasitoids has been observed in the 
laboratory (Pivnick 1993). Parasitized host 
larvae moved to the leaf litter and senes- 
cent leaves that become part of the litter 
where parasitoid pupation took place. 
This was confirmed in the field where co- 
coons successfully overwintered (Mason, 



86 



Journal of Hymenoptera Research 



unpublished). In both situations very few 
parasitoid cocoons were found on stand- 
ing plant parts that are harvested. 

Parasitism of M. configiimta by M. we- 
dintor clearly had a greater impact on the 
amount of food consumed by the host 
than parasitism by B. flavescens (Figs, lb 
and 2b). The results observed for M. me- 
diator are consistent with those observed 
for solitary koinobiont Microgastrinae 
Braconidae parasitoids attacking Lepidop- 
tera in other systems (Rahnian 1970, Sajap 
et al. 1978, Parkman and Shepard 1981, Ta- 
naka et al. 1984, Powell 1989). In contrast, 
Lepidopteran hosts parasitized by at least 
some gregarious koinobiont Braconidae 
and polyembryonic Encyrtidae species 
consume significantly more than non-par- 
asitized hosts and take longer to develop 
(Rahman 1970, Hunter and Stoner 1975, 
Gobbi et al. 1993, Byers et al. 1993). 

The decrease in feeding and weights af- 
ter a maximum peak is reached (Figs. 1 
and 2) is linked to cessation of feeding and 
gut evacuation associated with prepupal 
development in non-parasitized larvae, 
and probably to cessation of feeding by 
parasitized larvae which move to the leaf 
litter (M. mediator, see Pivnick 1993) or 
form earthen cells (B. flavescens, see Arthur 
and Mason 1985) where they complete de- 
velopment and pupate. The different 
times to peak weights and consumption 
between trials may be related to differenc- 
es in food quality, although the same plant 
variety, growing conditions, and leaf type 
were used in each trial. Genetic differenc- 
es between cohorts of M. coiifigurata and/ 
or the parasitoid species may also have 
caused this variation. 

The variability (i.e. large standard er- 
rors) observed, particularly on measure- 
ments made on day 13 or later (Figs. 1 anci 
2), may be associated with unexplained 
mortality during the experiments, differ- 
ences between sexes, or stochastic effects 
because of the relatively small numbers of 
experimental individuals (n ranged from 
5-9 and 7-9 for non-parasitized and par- 



asitized hosts in the B. flavescens trials and 
from 10 and 4-9 for non-parasitized and 
parasitized hosts in the M. tiiediator trials). 
The genetic diversity of individuals (even 
though hosts were from the same egg 
mass and parasitoids from the same fe- 
male) may have also been a factor. 

Natural controls, including parasitoids, 
have a major impact on bertha armyworm 
populations (Mason et al. 1998) and IPM 
programs should be developed around 
them. Chemical insecticides are used 
when 5"" and 6"' instar larvae are present 
on ripening canola pods (their feeding ac- 
counting for more than 86% of total larval 
consuniption). Wylie and Bucher (1977) 
reported that B. flavescens, like bertha ar- 
myworm, is univoltine. Although M. me- 
diator is bivoltine through most of its 
range in Eurasia where M. brassicae is also 
bivoltine, Johansen (1997) found it in a 
univoltine M. brassicae population in Nor- 
way (although Pivnick [1993] showed that 
M. mediator will diapause, further study is 
needed to determine if it would be uni- 
voltine on bertha armyworm). Adult B. 
flavescens and M. mediator are not likely to 
be affected by current chemical control 
practices because they attack early (1^' to 
3''') instar larvae and would disappear 
from the crop before 5"' and 6"' instar lar- 
vae are present. Even if M. mediator proves 
to be bivoltine in western Canada adults 
would probably move out of the canola 
crop seeking alternate hosts in adjacent 
habitats because no bertha armyworm in 
the appropriate stages would be available. 

Bertha armyworm larvae parasitized by 
B. flavescens feed until they reach the 6"' 
instar; thus the parasitoid would be neg- 
atively impacted by insecticides because 
the susceptible host is present in the crop 
being sprayed. Also, the parasitized larvae 
continue to feed (causing crop damage) 
during pod development and the benefits 
of parasitism by 6. flazK'scens are only ev- 
ident the following year. In contrast, M. 
confignrata larvae parasitized by M. medi- 
ator have ceased to feed (and consumed 



Volume 10, Number 1, 2001 



87 



600 
550 
500 

O) 450 

E 

'Z 400 

t 350 
^ 300 

^ 250 

LL 

c 200 

CD 


^ 150 

100 

50 





A 



Non-parasitized 
Parasitized 




DayO Day 4 Day 8 Day 1 1 Day 14 Day 19 

Days Post Parasitism 



Day 21 Day 24 



250 



.^ 200 



150 



100 



50 - 



Non-parasitized 
Parasitized 




Days 0-4 Days 4-8 Days 8-11 Days 11-14 Days 14-18 Days 18-21 Days 21-24 

Consumption Interval 

Fig. 1. Effect of parasitism (Trial #1) by B. flnvciicois on M. coiiftgiirata: a) Mean fresh weight (±SEM) of 
parasitized and non-parasitized Mnnwstya configurata larvae; b) Mean food consumption (±SEM)by parasitized 
and non-parasitized Miviicstm coiifigumtn larvae. 



<20% of the total consumption of non- 
parasitized larvae) by the time non-para- 
sitized 4"' instar larvae are present, before 
the switch to feeding on canola pods by 
later instar larvae. Further, M. mediator 
have formed cocoons and would likely be 



only minimally affected by insecticides 
used on 5"' and 6"' instar M. configurata, 
although this needs to be confirmed. 
Therefore, M. mediator appears to be a 
good candidate agent for use in the IPM 
of M. configurata because: 1) damage to the 



88 



Journal of Hymenoptera Research 



600 
550 - 
500 - 
450 
400 - 



5 350 - 
^ 300 - 

^ 250 - 

LL. 

^ 200 
(U 

^ 150 
100 - 
50 




Non-parasitized 
Parasitized 




Day Day 4 Day 8 



Day 11 Day 13 Day 15 Day 17 Day 19 
Days Post Parasitism 



Day 21 



0) 
■D 

E 



c 
o 
O 

c 

TO 
<D 



200 



150 



100 



50 



Non-parasitized 
Parasitized 



B 




Days 0-4 Days 4-8 Days 8-11 Days 1 1-13Days 13-15Days 15-17Days 17-19Days 19-21 

Consumption Interval 

Fig. 2. Effect of parasitism (Trial #1) by M. mediator on M. amfiguyaUv. a) Mean fresh weight (±SEM) of 
parasitized and non-parasitized Mnnicstm coufiguratn larvae; b) Mean food consumption (±SEM)by parasitized 
and non-parasitized Mamcstra coiifignrdtn larvae. 



host plant by parasitized larvae occurs agents such as chemical or biological in- 

when foliage is abundant and ceases be- secticides targeted at 4"' to 6"' instar bertha 

fore pod development begins; and 2) it armyworm which are applied after M. nie- 

should be compatible with other control diator has eliminated a portion of the pest 



Volume 10, Number 1, 2001 



population. Caged field release studies are 
needed to verify these hypotheses. 

The results obtained in this study sug- 
gest that for M. configiirnta parasitism by 
B. flavescens results in significantly de- 
creased consumption of food (and proba- 
bly less damage to canola crops) resulting 
in lower biomass production but does not 
reduce the length of time that the pest 
feeds in the crop and parasitism by the 
braconid M. mediator causes a significant 
reduction in the amount of food con- 
sumed and the length of time the host 
feeds compared to non-parasitized M. con- 
figurata. Parasitism and the findings pre- 
sented here should be taken into consid- 
eration when developing integrated pest 
management strategies for the bertha ar- 
myworm. 

ACKNOWLEDGEMENTS 

We thank Jennifer Prins for technical assistance. B. 
Thompson provided helpful statistical advice. L. 
Braun and U. Kuhlmann provided helpful comments 
of an earlier draft of the manuscript. 

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J. HYM. RES. 
Vol. 10(1), 2001, pp. 91-100 

Natural Alternative Hosts of Eulophidae 
(Hymenoptera: Chalcidoidea) Parasitoids of the Citrus Leafminer 
Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae) in the 

Mediterranean Basin 

Bruno Massa, Maria Concetta Rizzo, and Virgilio Caleca 

Istituto di Entomologia agraria, Universita degli Studi di Palermo, Viale delle Scienze 13, 
90128 Palermo, Italy. E-mail: zoolappl@vimpa.it 



Abstract. — The entomofauna linked to native flora in and around citrus groves was studied in 
Italy and Jordan in order to find alternative hosts of eulophid parasitoids of the Citrus Leafminer 
(CLM), Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae). Twenty new associations (12 in 
Italy, 8 in Jordan) among native and exotic CLM parasitoids and leafminers were found. Two 
new alternative hosts were recorded for Citrostichus piln/llociiistoides (Narayanan) (an unidentified 
Nepticulidae on Pistacia lentiscus L. and Stigmella sp. on Rubus ulmifolius Schott, in Sicily and 
Jordan respectively) and 1 for Cirrospiliis iiigenuus Gahan (Agromyzidae on Salix sp., in Jordan). 
Five new alternative hosts were recorded for Semielacher petiolatus (Girault) (in Sicily Liriomyza sp. 
on Mcrciirialis annua L., Chromatomyia horticola (Goureau) on Sonchiis spp., Cosvioptcrix pulchrimella 
Chambers on Parictaria diffusa M. et K., and Stigmella aurcUa (Fabr.) on Rubus ulmifolius Schott; in 
Jordan, Dialcctica scalariella Zeller on Echium sp.). The other 12 new associations of CLM parasitoids 
with leafminers found in both countries include NeochrysocJiaris formosa (Westwood) (4 new hosts), 
Cirrospilus variegatus (Masi) (5 new hosts), Ratzeburgiola incompleta Boucek (1 new host), Rntzcbur- 
giola cristata (Ratzeburg) (1 new host), and Asccodcs dclucchii (Boucek) (1 new host). Data reported 
here suggest that native vegetation harbours alternative hosts for both native and exohc parasit- 
oids. They also underline that more attention should be paid to the understanding of ecology and 
biology of parasitoid species in order to use appropriate exotic enemies in biological control, 
preserving at the same time indigenous parasitoid communities. 



The Citrus Leafminer (CLM), Phyllocnis- CLM in many countries and in some case 

tis citrella Stainton (Lepidoptera: Gracillar- also have been recovered in neighbouring 

iidae), is presently considered to be a se- countries (Schauff et al. 1998). 

rious threat to young citrus trees in the Native plants are an important source of 

Mediterranean region and other countries, biological diversity in agroecosystems and 

w^here it has expanded its range in the last are known to harbour natural enemies of 

seven years. The CLM attacks new leaves phytophagous pests of cultivated plants, 

causing a loss of photosynthetic area. Seni- supplying alternative food, refuges and 

ielacher petiolatus (Girault), Citrostichus hosts (McMurtry and Johnson 1965, Pow- 

phyllociiistoides (Narayanan) and Cirrospi- ell 1986, Altieri 1991, Ragusa Di Chiara 

lus ingenuus Gahan (Hymenoptera: Eulo- 1991). They provide a diverse source of 

phidae) are considered among dominant food for many species of polyphagous nat- 

natural enemies of CLM in its original ural enemies, which in turn may parasit- 

range (India and South Asia) and in Aus- ize phytophagous insects of cultivated 

tralia (Smith et al 1997, Schauff et al. 1998). plants in seasons when they are abundant. 

Eulophid parasitoids have been selected Studies of phytophagous insects are of- 

for biological control programmes against ten directed at species attacking cultivated 



92 



Journal of Hymenoptera Research 



plants and, less so, species feeding on na- 
tive ones; thus our knowledge of the hosts 
of parasitoids on cultivated plants is ex- 
tensive, whereas we have only scattered 
data on the alternative hosts available to 
these parasitoids on native plants. Preser- 
vation of these reservoirs of antagonists 
may prove valuable when parasitoids uti- 
lize hosts that are not pests of cultivated 
plants, and when biological control de- 
pends on multiple natural enemy species, 
as in the case of the CLM. 

The present study is part of a research 
project examining the entomofauna of na- 
tive flora carried out in 11 citrus orchards 
of Sicily (Italy), the results of which were 
partly published (Caleca et al. 1997, Mineo 
et al. 1997a, 1997b, Caleca 1998, Caleca et 
al. 1998, Mineo and Sinacori 1998, Rizzo et 
al. 1999, Massa and Rizzo 2000). Some 
new findings are presented that highlight 
interesting relationships between CLM 
parasitoids and their non-pest hosts ex- 
ploiting native plants. 

MATERIALS AND METHODS 

Native floras associated with citrus 
groves in Sicily (Italy) have been studied 
in detail by Raimondo et al. (1979); they 
amount to about 200 species involving 
mainly herbs and shrubs. During the four 
years of our research project (1997-2000), 
we collected about 40 of the most common 
species belonging to this flora, and about 
10 belonging to riverine flora sometimes 
occurring in the neighbouring areas of cit- 
rus orchards in Sicily. About 250 g of each 
plant species were collected monthly 
along at least two perpendicular transects 
inside 11 citrus groves and along their pe- 
rimeter. Leaves infested by miners were 
placed in Petri dishes with wet paper at 
25° C, 65% r.h. and L14:D10. All phytoph- 
agous species and parasitoids that 
emerged were mounted and identified. 
Further samples were gathered by the se- 
nior author during a research trip to Jor- 
dan between 21 and 29 May 1999 in the 



following localities: Al Bahhath (Amman), 
Aqaba and Dana Village. 

RESULTS 

In Sicily, 40 host-parasitoid associations 
involving phytophages of native plants 
and antagonists of CLM were already 
known (Caleca et al. 1997, Mineo et al. 
1997a, 1997b, Caleca 1998, Caleca et al. 

1998, Mineo and Sinacori 1998, Rizzo et al. 

1999, Massa and Rizzo 2000). They are list- 
ed in Fig. 1 together with the 12 (Italy) and 
8 (Jordan) associations, previously unno- 
ticed, recorded in the present paper. Data 
for new records are reported in Table 1. 

The parasitoids we found on indige- 
nous leafminers belong to two quite dif- 
ferent kinds: exotic CLM biological control 
agents (i.e., Semielacher petiolatus, Citrosti- 
chiis phyllocnistoides and Cirrospilus iiigen- 
uus), which possibly have switched over 
onto indigenous hosts after their introduc- 
tion or immigration, and native parasit- 
oids, which in turn have switched over 
onto the invading CLM. Among the latter, 
some (i.e., Neocluysocharis formosa (West- 
wood), Ratzeburgiola iucompleta Boucek 
and P}iigalio soeDiius (Walker)) are quite 
common on indigenous hosts in Sicily 
(Fig. la), while three of them (i.e., Digly- 
phiis isaea (Walker), Ratzeburgiola cristata 
(Ratzeburg) and Cirrospilus variegatus 
(Masi)) have not yet been recorded on 
CLM in the island, although they have 
been reported on this host in other coun- 
tries (Schauff et al. 1998). It should be 
pointed out that, even if their parasitiza- 
tion does not always reach relevant val- 
ues, most CLM parasitoids engage in host- 
feeding which contributes additional mor- 
tality. For example in Algeria Guenaoui 
and Dahliz (1997) attributed as much as 
20-50% of CLM larval mortality to host- 
feeding and in Sicily it may approach 15% 
(pers. obs.). 

Among the eight new host-parasitoid as- 
sociations reported from Jordan (Fig. lb), 
five concern eulophids (S. petiolatus, N. for- 
fuosa, C. phyllocnistoides and C. iugeiiuus) 



Volume 10, Number I, 2001 



93 



Plants 



Phytophagous insects 



Eulophids known to attack 
Phvllocnislis cilrella 



a) Sicily 



Sonchiis ( 3 spp ) 


4l" 


Chromalomyia horlicola (Goreau) 






Neochnsocharis/ormosa (West wood)'" | 


Reichardia picroides (L ) 




Uriica (2 spp ) 


Agrom\za hwmalis Becker 


Chnsocharis peniheus (Walker) | 




Mercunalis annua L 


Lir,om,-a sp 






Beta vulgaris L 


Scaplonn-a pallida (Fallen) 


Dvgliphus isaea (Walker)' [ 


♦ 

< 
< 


Snnrmum (2 spp ) 


Euleia heraclei ( L ) 




Borago officinalis L 


Dialeclica scalariella Zeller 


Semielacher petiolaius (Girault) | 


Ainplex rosea L 


Chnsoesthia drurella ( F ) 




Chenopodium album L 


Chnsoeslhia sexgiillella (Thunberg) 


Raizeburgiola incompleta Boucek | 


Panefaria ( 2 spp ) 


Cosmoplerix pulclinmella Chambers 




"* 


l.alhvrus pralensis L 


Liriom\-a congesta (Becker) 


Ralzeburgiola crislala (Ralzeburg)' | 


< 

< - 


*,-r' J.--;..--'..--' '•-< 
^-'/'-■■■".-■■'' 

fe-''' 


■ 


Ruhiis ulmifolius Schott 


Stigmella aurella (F ) 




Olea europaea L var 
svlvestns Brot 


Baclrocera oleae (Gmel ) 


Pmgalio soemius (Walker)'"' | 






Cellis auslralis L 


PMIonon'Cler milleriella (Staudinger) 


Pnigalio agraules (Walker) | 


* 


Pislacia lenliscus L 


unidentified Nepticulidae 




1 ihiirniim linus L 


PInllononcier lanlhanella ( H ubner ) 


Cirrospilus variegalus (Masi)* | 


* - 


Crataegus monog\-na Jacq 


Leucopiera malifoliella (Costa) 




Ulmus sp 


Sligmella sp 


Cirrospitus tvnciis Walker | 




PInllononcier conlifoliella ( Schrank ) 




5o/« pedicellala Desf 


PMIononcier duhilella ( H -S ) 


Cirrospilus diallus Walker ] 




Stigmella sp 




So/« o/Ao L 


PMIocnistis saligna (Zeller) 


Cirrospilus pictus ( Nees ) 




Japanagromvza salicifolii (Collin ) 






PInllocnislis lahvrinlhella ( Bjerk ) 


Cilrosuchus phyllocnisloides ( Narayanan ) 


Populus nigra L 


PInllocnisiis unipunciella (Stainton) 






Caloplilia sligmatella i F ) 


Apotetrastichus poslmargmalis ( Boucek ) 




Sligmella Irimaciilella ( Haworth) 






Messa horlulana klug 




b) Jordan 






Echnim sp 


Dialeclica scalariella 


Semielacher peliolalus 






Mallhtolo incana ( L ) 


Liriom\^a hryonioe { Kaltenbach ) 


Neochrx'socharis formosa 






Chrozophora tinclona ( L ) 


^ 


unidentified Nepticulidae 


<— 


Cirrospilus variegalus * * 


Ri/Ai« ulmifolius 


Sligmella sp 


Cilrosuchus phyllocnisloides 


\^--" 








Ralzeburgiola incomplela" 


Salix sp 


unidentified Agromyzidae 


)**=iiiiiir^ 




Asecodes delucchii ( Boucek ) * * 




Cirrospilus ingenuus Gahan 



< — New records 

<— Known records, parasitoid-host relationships after Calecaet al W')7. Caleca |W8, Caleca& Lo Verde IW8. Caleca el al IQ98. Mineoel al 1997a, b. 

Mineo& Sinacori 1998, Rizzo el al 1999. Massa& RizzoCOOO) 
* Species recorded in other Medilerranean areas on P- cilrella. but unrecorded on Ihis hosi in Sicily 
** Species recorded in other Medilerranean areas on P cilrella, hul unrecorded on this host in Jordan 
'" This species was recorded by Caleca el al ( 1997) on Beia vulgaris ex Scapionnza sp , after collecting new malenal this host has been idenlilled as 

Scaptomyza pallida 
''' This species was recorded by Caleca ( 1998) on Salix alba e\ unidenlified Agromy/idae. after collecting new material Ihis host has been identified as 
Japanogromyza salicifolii 

Fig. 1. Associations between eulophid parasitoids of Phyllocnistis citirlla and other phytophagous insects ex- 
ploiting native plants in citrus groves in Sicily, Italy (a) and Jordan (b). 



known to attack CLM in the country 
(Schauff et al. 1998, Mineo 1999). C. varie- 
gatus, R. incotnpleta and A. delucchii (Bou- 
cek) have been recorded as CLM parasit- 
oids in other countries of the Mediterra- 
nean Basin, but until now have been 
found only on native plants in Jordan, al- 
though this may be due to the paucity of 
information on CLM in this country. Some 
details on the new findings are reported 
below. 

SeniielacJier pu'tiolntus (Girault) 

Originally described from Australia, S. 
petiolatus has also been reported from the 



Solomon Islands (Boucek 1988, Schauff et 
al. 1998). It was introduced to Oman, Sy- 
ria, Israel, Egypt, Cyprus, Greece, Turkey, 
Tunisia and Morocco (Michelakis 1997, 
Argov and Rossler 1996, FAO 1996, Nia et 
al. 1997, Rossler and Argov 1997, Hamed 
et al. 1999); it spread spontaneously in It- 
aly, Algeria and Jordan (Mineo et al. 1998, 
Schauff et al. 1998, Mineo 1999). Before be- 
ing reared from Agronn/za hiemalis Becker 
(Diptera: Agromyzidae) (Massa and Rizzo 
2000), the only host previously known for 
this species was the CLM (Schauff et al. 
1998), and, from 1998 onwards, it became 
one of the most important parasitoids of 



94 



Journal of Hymenopthra Research 



this pest in Sicily (Caleca et al. 1998, Mineo 
and Mineo 1999a). The five new hosts hst- 
ed in this paper (Table 1), comprising 
three Lepidoptera (Cosmopterigidae, Nep- 
ticulidae and Gracillariidae) and two Dip- 
tera (Agromyzidae), are widespread in the 
Mediterranean region. 

It should be noted that this species ap- 
peared in Sicily on Chromatomyia horticola 
(Goureau) (Diptera: Agromyzidae) about 
one year after its release in 1996 in Tunisia 
(FAO 1996). The availability of alternative 
hosts that provide refuge and food for S. 
petioJatus during seasons of low CLM pop- 
ulation density, could partly explain the 
quick spread and establishment of this 
species, both in countries where it has 
been released and in neighbouring sites 
(Caleca et al. 1998, Schauff et al. 1998, Mi- 
neo and Mineo 1999a). 

Citrostichus phyllociiistoides (Narayanan) 

This species is known from Afghani- 
stan, China, India, Indonesia, Japan, 
Oman, Pakistan, Taiwan, Thailand, 
South Africa, Sudan, Swaziland (Boucek 
1988, FAO 1996, Schauff et al. 1998), and 
has been introduced to Cyprus, Greece 
and Italy (Sicily) (Michelakis 1997, FAO 
1996, Mineo and Mineo 1999b), and Aus- 
tralia and Israel (where it is not consid- 
ered established) (Smith et al. 1997, Ar- 
gov and Rossler 1996). It probably 
spread to Jordan from Israel (Mineo 
1999). Although recorded as a parasitoid 
of the CLM (Boucek 1988, Ujiye and 
Adachi 1995, Wu and Lin 1998), it also 
has been reported to parasitize the 
nymphs of Trioza obsoleta Buckton (Ho- 
moptera: Psyllidae) a gall former on Di- 
ospijros nielanoxylon (Roxb.) (Dash and 
Das 1997). Our records concern two ad- 
ditional Lepidoptera (Table 1); according 
to Nieukerken (pers. comm.) the mines 
on Rubus ulmifolius Schott found in Jor- 
dan belong to a Stii^tfiella species (Nep- 
ticulidae), possibly Stigmella aurella 
(Fabr.), a previously unrecorded host for 
this parasitoid. 



Cirrospilus ingenuus Gahan 

Also known as its synonym C. quadris- 
triatiis (Subba Rao and Ramamani), this 
species has been recorded from Austra- 
lia, China, India, Indonesia, Japan, Ma- 
laysia, Oman, Taiwan, Thailand (Smith 
et al. 1997, Schauff et al. 1998), and intro- 
duced to Cyprus, Turkey, Israel, Syria, 
Egypt, Tunisia, Morocco, Florida and 
Mexico (Argov and Rossler 1996, FAO 
1996, Perales-Gutierrez et al. 1996, Ha- 
med et al. 1999, LaSalle et al. 1999). It has 
also spread to Jordan and North Egypt, 
probably from other countries of the 
Mediterranean Basin (Schauff et al. 1998). 
It is generally considered to be a domi- 
nant parasitoid of P. citrella (e.g., Thai- 
land, Taiwan and Japan: Ujiye et al. 
1996), but also has been recorded as a 
parasitoid of Rhynchaeiuis maugiferae 
Marshall (Coleoptera: Curculionidae) in 
India (Peter and Balasubramanian 1984). 
Agromyzidae previously has not been 
recorded as a host for this parasitoid. 

Cirrospilus variegatus (Masi) 

Also known as its synonym Zagrammo- 
soffia variegatum, it occurs in the Mediter- 
ranean Region, North and East Africa, 
Central and South Asia, West Indies (Bar- 
bados), Australia and New Zealand (Bou- 
cek 1988; Yefremova 1996). This species 
was described from Italy by Masi (1907), 
as parasitoid of Metriochroa latifoliella (Mil- 
liere) (Lepidoptera: Gracillariidae). It is 
also known to parasitize many species of 
small leaf-mining Lepidoptera (Boucek 
1988, Yefremova 1996) and the CLM in 
Libya, Spain and Turkey (Schauff et al. 
1998). During this study it was found on 
two Nepticulidae leafminers and on three 
Diptera (one Tephritidae and two Agro- 
myzidae) (Table 1), all previously unre- 
cognised as hosts. 

Ratzeburgiola incompleta Boucek 

Recorded from central Europe and 
many countries of the Mediterranean Ba- 



Volume 10, Number 1, 2001 



95 



Table 1. List of new host records for Eulophidae emerged from leafminers reared from native plants 
collected in Italy and Jordan. 



l.ulophid 



I lost spri ir 



Host plant 



Sciiiiclachcr pctiolntu 
(Girault) 



ides (Narayanan) 



Cirrospilus iiigcniiiis 

Gahan 
Cirrospiliis varicgntiis 

(Masi) 



Ratzcburgioln iiicoiiiplcta 

Boucek 
Ratzcbiirgioln cristata 

(Ratzeburg) 
Neochrx/socharis foiiiwsn 

(Westwood ) 



Asccodcs dclucchii 
(Boucek) 



Chnvimtoim/in horticoln 
(Goureau) (Diptera: 
Agromyzidae) 

Lirioiin/za sp. (Diptera: 
Agromyzidae) 

Cosiiioptcrix piilchriincl- 
la Chambers (Lepi- 
doptera: Cosmopter- 
igidae) 

Stiginclla nurclla {Fnbr.) 
(Lepidoptera; Nepti- 
culidae) 

Dialectica scaliviclln 
Zeller (Lepidoptera: 
Gracillariidae) 

Lepidoptera: Nepticu- 
lidae 

StigiucIIa sp. (Lepidop- 
tera: Nepticulidae) 

Diptera: Agromyzidae 

C. horticoln 

Lirionn/za coiigcsta 
(Becker) (Diptera: 
Agromyzidae) 

Eulcia hcradci (L.) 
(Diptera: Tephriti- 
dae) 

Lepidoptera: Nepticu- 
lidae 

Lepidoptera: Nepticu- 
lidae 

Diptera: Agromyzidae 

L. coiigcstn 

D. scalariclla 

Lt'ticoptcrn nialifoliclln 
(Costa) (Lepidop- 
tera: Gracillariidae) 

Lirionn/zti bn/oninc 
(Kaltenbach) 

Lepidoptera: Nepticu- 
lidae 

l~>iptera: Agromyzidae 



Soiichiis spp. 



Mcrcurialis annua L. 

Parictaria diffusa M. 
and K. 



Rubus uhiufolius Schott 

Echiuni sp. 

Pistacm Iciitiscus L. 

R. uhiiifolius 

Salix sp. 

Rcichardia picroidcs (L.) 

Lathyrus pratcnsis L. 

Snn/rniuin pcrfoUatuni 
(L.) 

P. Icutiscus 

Chrozophora ti}U'toria 

(L.) 
Salix sp. 

L. pratcnsis 

Borago officinalis L. 

Crataegus inonogifna 
Jacq. 

Mattlnola uicana (L.) 

C. tinctoria 

Salix sp. 



Ital}, Parco d'Orleans 26 
(Palermo) 3.V.97, 
21.in.99 

Italy, Borgo Molara 1 9 

(Palermo) 29.XI.98 

Italy, Zucco (Palermo) 49 
6.VI.00, Borgo Mo- 
lara 26.1.99, 14.III.99 

Borgo Molara 1 9 

17.VIII.99, Parco 

d'Orleans 20.IX.99 
Jordan, Dana Village 1 9 

25.V.99 



Zucco 4.IV.00 


1(5, 


19 


Jordan, A! Bahhath 


16, 


19 


(Amman) 23.V.99 






Al Bahhath 23.V.99 


19 





Italy, Collesano (Paler- 1 ' 

mo) 20.V.99 
Collesano 20.V.99 1 • 



Italy, Petralia Sottana 3 9 
(Palermo) 6.VI.99 

Zucco 27.VIII.99 1 6 

Jordan, Aqaba 27.V.99 36,49 

Al Bahhath 23.V.99 \6 

Collesano 20. V.99 29 

Italy, Bagheria (Paler- 1 9 

mo) 20.XI.97 

Zucco 7.VIII.99 79 

Al Bahhath 23.V.99 Id, 29 

Aqaba 27. V.99 2d, 19 

Al Bahhath 23.V.9q 1 9 



96 



Journal of Hymenoptera Research 



sin (Boucek 1969, 1970, Schauff et al. 1998), 
this species is known to parasitize Holo- 
cacista rivillei Stainton (Lepidoptera: He- 
liozelidae) (Boucek, 1969), Phyllouonjcter 
corylifoUclla Hiibner (Lepidoptera: Gracil- 
lariidae) (Mineo and Sinacori 1998), Cos- 
mopterix pulchrimella Chambers (Lepidop- 
tera: Cosmopterigidae) (Rizzo and Mineo 
1997), Liriomyza trifolii (Burgess) (Diptera: 
Agromyzidae) (Freidberg and Gijswijt 
1983), Urioviyza sp. (Diptera: Agromyzi- 
dae) (Rizzo and Mineo 1997), and Agro- 
myza hiennilis (Massa and Rizzo 2000). It is 
also recorded as CLM parasitoid (Azawi 
1997, Schauff et al. 1998), and as the most 
abundant native CLM parasitoid in Tur- 
key (Uygun et al. 1997) and Israel (Rossler 
and Argov 1997). 

Ratzeburgiola cristata (Ratzeburg) 

Known from the whole Europe (Boucek 
and Askew 1968, Rizzo and Mineo 1997, 
Schauff et al. 1998) parasitizing Plnjllono- 
rycter nigrescentella Logan (Lepidoptera: 
Gracillariidae) and Cosmia trapezina L. 
(Lepidoptera: Noctuidae) (Boucek and 
Askew 1968), Chrysoesthia sexguttella 
(Thunberg) (Lepidoptera: Gelechiidae), 
Cosuiopterix pulchrimella and Stigjiiella aii- 
rella (Rizzo and Mineo 1997), and the CLM 
in Spain (Schauff et al. 1998). Agromyzidae 
previously has not been recorded as a host 
for this parasitoid. 

Neochrysocharis foiiiiosa (Westwood) 

This species is known from the Palearc- 
tic, Asia and Africa (Boucek and Askew 
1968). It develops as a primary endopar- 
asitoid of larvae, and rarely eggs, of leaf- 
miners (Hansson 1990), and is known as 
parasitoid of P. citrella in Cyprus, Greece, 
Israel, Italy, Japan, Jordan, Spain, Tunisia 
and Turkey (Caleca et al. 1996, FAO 1996, 
Ujiye et al. 1996, Schauff et al. 1998). 

Asecodes delucchii (Boucek) 

Asecodes delucchii (= Teleopterus deluc- 
chii) is known throughout the Palearctic 
Region from England to Japan (J. LaSalle, 



pers. comm.), but has not been previously 
recorded in Jordan. It is known to attack 
P. citrella in Italy and Japan (Ujiye and 
Adachi 1995, Ujiye et al. 1996, Mineo 
1999), and has also been recorded as a par- 
asitoid of Caliroa cerasi L. (Hymenoptera: 
Tenthredinidae) and Phyllonorycter messan- 
iella (Zeller) (Lepidoptera: Gracillariidae) 
(J. LaSalle, pers. comm.). Agromyzidae 
previously has not been recorded as a host 
for this parasitoid. 

DISCUSSION 

The introduction of exotic polyphagous 
parasitoids could decrease native parasit- 
oids competing for the same food resource 
(Bennett 1993, Duan et al. 1996); thus, due 
to the naturally low density of their pop- 
ulations, native polyphagous parasitoids 
may undergo dramatic decrease to be- 
come locally extinct (LaSalle 1993). For 
these reasons, to find the best control 
agent of a noxious insect before using it in 
biological control programs LaSalle (1993) 
suggested carrying out research on the bi- 
ology and ecology of species of parasit- 
oids that are considered antagonists of the 
host. 

Since 1993, when P. citrella colonised 
Mediterranean citrus groves, endemic bi- 
ological diversity represented the poten- 
tial resource for biological control (cf. 
LaSalle 1993). A dozen native polypha- 
gous eulophids were found to parasite it. 
These species constituted the parasitoid 
community living on leaf-miners of the 
native flora. Research carried out in Sicily 
listed a total of 47 associations involving 
native eulophids that parasitize the CLM 
(Caleca et al. 1997, Mineo et al. 1997a, 
1997b, Caleca 1998, Caleca et al. 1998, Mi- 
neo and Sinacori 1998, Rizzo et al. 1999, 
Massa and Rizzo 2000, present study) (cf. 
Fig. la). Among CLM parasitoids, the ge- 
nus Cirrospilus (Hymenoptera: Eulophi- 
dae) played a dominant role, particularly 
C. pictus (Nees) in Sicily, Algeria and 
Spain (Caleca et al. 1998, Guenaoui and 
Dahlis 1997, Vercher et al. 1997). Due to 



Volume 10, Number 1, 2001 



97 



the spread of CLM, many researchers 
planned the introduction of its specific 
control agents; in the Mediterranean Basin 
today at least 6 exotic species, known as 
dominant parasitoids of P. citrella, have 
been introduced (Argov and Rossler 
1996). Among them S. petiolntus sponta- 
neously colonised Sicily (Mineo et al. 
1998), probably from N Africa, where it 
had been introduced, while Ageniaspiis ci- 
tricola Logvinovskaya (Hymenoptera: En- 
cyrtidae) and the eulophids Qundrnstichus 
sp. and C. pJn/Uociiistoides were here ac- 
tively introduced (Siscaro et al. 1997, Mi- 
neo and Mineo 1999b). 

Our research led us to find 5 alternative 
new native hosts of S. petiolntus and 2 of 
C. phyllocnistoides, as well as another new 
host of C. ingenuus, eulophids previously 
known as dominant or specialist CLM 
parasitoids. We believe that alternative 
hosts, leaf-miners of native flora, contrib- 
uted to acclimatation of S. petiolntus and 
C. phyllocnistoides in Sicily, providing al- 
ternative food and shelter, mainly in win- 
ter and spring, when CLM populations 
decrease very much (Massa and Rizzo in 
press). As regards the interference deter- 
mined on native CLM parasitoids by ex- 
otic ones, S. petiolntus in 1998 represented 
as much as 38% of all the parasitoids in 
an orange grove, with an average parasit- 
ization rate of 6.9%, peaking to 87.5% in 
September (Caleca et nl. 1998), while in 
1999 it represented 89% of all the parasit- 
oids in a lemon grove, with a peak of par- 
asitization rate of 69.6% (Mineo and Mi- 
neo 1999b), playing a dominant role in the 
CLM control. C. pJn/llocfiistoides on the 
contrary seems to be still sporadic in Sicily 
(Mineo and Mineo 1999b). 

Among native CLM parasitoids in Sici- 
ly, in 1998 C. pictus reached 7.9% of par- 
asitization rate, while all the other para- 
sitoids did not exceed 2% (Caleca et nl. 
1998), values already known before the in- 
troduction of exotic parasitoids. As re- 
gards the parasitization on hosts of native 
flora, even if our data do not present a 



significant quantitative analysis, from the 
qualitative point of view it seems that the 
community structure of parasitoids did 
not change after the introduction of exotic 
species (Caleca et nl. 1997, Mineo et nl. 
1997a, 1997b, Caleca 1998, Caleca et nl. 

1998, Mineo and Sinacori 1998, Rizzo et nl. 

1999, Massa and Rizzo 2000). The number 
of individuals of S. petiolntus and C. phyl- 
locnistoides found parasitizing native hosts 
indeed is still low, compared with the 
whole number of parasitoids (564) ob- 
tained during our research (Massa and 
Rizzo in press). 

Finally, our results point out that native 
flora within and at the edge of citrus 
groves strengthens the biological control 
of P. citrelln, providing alternative hosts to 
its parasitoids, native as well exotic, main- 
ly in the winter-spring seasons when P. 
citrelln density is very low. Additionally, 
they stress the importance of knowledge 
of parasitoid biology and ecology to opti- 
mise their use in biological control pro- 
grams, as well the conservation of native 
parasitoid communities. 

ACKNOWLEDGMENTS 

The authors thank Prof. Pietro Mazzola and Dr. 
Lorenzo Gianguzzi (Dipartimento di Scienze Botani- 
che, Palermo) for plant identification; Prof. Luciano 
Siiss (Istituto di Entomologia agraria, Milan), Dr. Pa- 
olo Triberti (Museo di Storia Naturale, Verona), Dr. 
Erik Van Nieukerken (Rijksmuseum van Naturlijke 
Historic, Leiden) and Dr. John LaSalle (CABI Biosci- 
ence UK Centre, Biology Dept., Ascot) for the iden- 
tification of Agromyzidae, Gracillariidae, Nepticuli- 
dae and some Eulophidae, respectively; Prof. Ahmad 
Katbeh-Bader (Horticulture and Plant Protection 
Dept., Jordan Univ., Amman), Dr. Salvatore Blando, 
Rocco Lo Duca, Dr. Gabriella Lo Verde and Dr. Val- 
entina Lo Verde (Istituto di Entomologia agraria, Pa- 
lermo Univ.) for their help in the field; and Prof. At- 
tilio Carapezza (Palermo University) for improving 
English. Paper funded by C.N.R. and M.U.R.S.T. 

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Schauff, M. E., J. LaSalle, and G. A. Wijesekara. 1998. 
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Ujiye, T. and I. Adachi. 1995. Parasitoids of the citrus 
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Journal of Hymenoptera Research 



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J. HYM. RES. 
Vol. 10(1), 2001, pp. 101-111 

Interactions Between Adults of Some Species of Netelia Gray 

(Hymenoptera: Ichneumonidae: Tryphoninae) and 

Their Caterpillar Hosts (Lepidoptera) 

Mark R. Shaw 
National Museums of Scotland, Chambers Street, Edinburgh, EHl IJF, UK 



Abstract. — Five species from three subgenera of Netelia were observed ovipositing on their hosts 
under laboratory conditions. Two species of the subgenus Bessobates oviposited without a separate 
insertion of the ovipositor beforehand. One species of the subgenus Netelia and two of the sub- 
genus Paropheltes employed a venom causing weak (often incomplete) temporary paralysis, ad- 
ministered as a separate action prior to oviposition. The venoms had no long-term adverse effect 
on hosts and it appears that these taxa, and perhaps all other groups of ectoparasitic koinobiont 
Ichneumonidae, lack the host-regulating venom systems that have been regarded as instrumental 
in the evolution of endoparasitism in some koinobiont cyclostome Braconidae. In all species sub- 
stantial parasitoid development was delayed until the final instar host had constructed its pu- 
pation site, wherein it was overwhelmed as a prepupa. Species of all three subgenera dumped 
eggs in the absence of hosts, and (if kept humid) such eggs subsequently split to reveal a living 
first instar larva (but not investigated in the subgenus Paropheltes). Species of all three subgenera 
indulged in non-destructive concurrent host-feeding, despite being supplied with honey solution 
ad libitum, making the necessary wounds on the host's body with their mandibles. Occasionally 
non-destructive host-feeding occurred on hosts that were not also being parasitised. 



The main purpose of this study was to 
investigate the effect of the venom of spe- 
cies of Netelia Gray (= Pniiiscus auctt. nee 
Schrank) (Ichneumonidae: Tryphoninae) 
on their caterpillar hosts, but in the course 
of conducting experiments observations 
were also made on host-feeding, oviposi- 
tion behaviour, egg dumping, egg hatch- 
ing and (in one species) copulation. 

Many koinobiont parasitic wasps — that 
is, those in which the host continues to de- 
velop after being parasitised — influence 
the subsequent physiology, behaviour and 
development of their hosts in various and 
often profound ways (Vinson and 
Iwantsch 1980, Beckage 1985, 1991, 
Lawrence 1986, Barnard and Behnke 1990; 
see also Quicke 1997). In some, injections 
of materials ("venoms" in the broadest 
sense) other than eggs by the female par- 
asitoid during the oviposition sequence 
are known to be at least partly responsible 



(Shaw 1981, Tanaka 1987, Jones 1987, Do- 
ver et al. 1988, Leluk and Jones 1989). In 
endoparasitoids the effect on the host of 
venoms independent of parasitisation as 
such is usually difficult to investigate di- 
rectly because the egg is injected concur- 
rently; that is, during a single insertion of 
the ovipositor. In some koinobiont cyclo- 
stome Braconidae, however, venoms ini- 
tially causing paralysis that is only tem- 
porary are injected as a preliminary and 
separate action prior to oviposition, and in 
some taxa the venom has been shown to 
switch the host, which initially recovers to 
resume feeding, to a prepupal stage in 
which it is arrested (Shaw 1981, 1983). 
This includes cocoon formation and the 
secretion of pupal cuticle, both of which 
happen precociously (i.e. an instar early) 
if the host is attacked in its nominally pen- 
ultimate larval instar, and two koinobiont 
cyclostome braconid genera, the ectopar- 



102 



Journal of Hymenoptera Research 



asitic RhysipoUs Foerster (Rhysipolinae) 
and the endoparasitic CUnocentrus Haliday 
(Rogadinae), were found to employ ven- 
oms having identical effects on their hosts. 
Although these taxa are not regarded as 
extremely closely related, the suggestion 
(Shaw 1983) that on at least one occasion 
the evolution of endoparasitism from ec- 
toparasitoid ancestry within the Exotheci- 
nae-Rhysipolinae-Rogadinae group of 
subfamilies was enabled as a result of 
these controlling venom systems, which 
arose originally as adaptations to the 
host's feeding and pupation biology, has 
been supported by other evidence (Whit- 
field 1992). 

Gauld (1988) envisaged that within the 
Ichneumonidae idiobiont endoparasitism 
arose from idiobiont ectoparasitic ances- 
try, and koinobiont endoparasitism fron"i 
koinobiont ectoparasitic ancestry. The 
mechanisms for such transitions remain 
uncertain but it is possible in principal 
that the latter route may have involved 
venom systems similar to those seen in 
some cyclostome Braconidae. However, 
evidence for such venom effects in Ichneu- 
monidae has not been directly sought, al- 
though it is known that in some endopar- 
asitoid subfamilies (particularly Campo- 
pleginae) complex physiological processes 
ensue from substances injected simulta- 
neously with eggs. Three extant groups of 
koinobiont ectoparasitoid Ichneumonidae 
are known: Adelognathinae (most inves- 
tigated species, parasitoids of sawfly lar- 
vae); the Poh/sphitictn Gravenhorst genus- 
group (= Polysphinctini sensu Townes 
1969, but see Wahl and Gauld 1998) of the 
Pimplinae (parasitising spiders); and Try- 
phoninae (parasitising sawfly and Lepi- 
doptera larvae). At least some members of 
each group are known to inject venoms 
causing temporary paralysis as separate 
actions prior to oviposition (as in Rln/si- 
polis and the endoparasitic CUnocentrus), 
offering particularly convenient opportu- 
nities to interrupt the oviposition se- 
quence so as to observe any venom effects 



in isolation. Indications exist (see Discus- 
sion) that in Adelognathinae and the Po- 
h/sphijjctn genus-group these venom sys- 
tems are innocuous, in the sense of caus- 
ing only temporary paralysis rather than 
exerting lasting control over the host's de- 
velopment, but for the third group, Try- 
phoninae, essential information is lacking. 
Because Tryphoninae generally kill the 
host as a prepupa, the possibility that they 
employ venoms that disrupt processes 
normally under endocrine control (as with 
RJiysipolis and Clinocoitrus) seemed partic- 
ularly worth examining. 

Tryphonines are predominantly (at least 
at the generic level) parasitic on Symphyta 
but the family contains the tribe Phytodie- 
tini, including the genera Netelia and Phy- 
todietus Gravenhorst, which attack Lepi- 
doptera larvae. In view of the ease of cul- 
turing their hosts, species of Netelia were 
chosen for study. Netelia species anchor 
their highly characteristic stalked black, 
and typically glossy, eggs externally on 
the thoracic segments of active exposed 
Lepidoptera larvae that are normally con- 
sumed only after constructing their pu- 
pation chambers (i.e. as prepupae). The 
genus contains species for which tempo- 
rary paralysis of the host has been report- 
ed as well as others that apparently ovi- 
posit without having this effect (e.g. Sten- 
ton 1910, Shevyrev 1912, Vance 1927). 
While many observations have been pub- 
lished on the egg structure and larval de- 
velopment of Netelia (e.g. Stenton 1910, 
Cushman 1913, Strickland 1923, Vance 
1927, Guppy 1961, Danks et al. 1979, Ellis 
1998), the possibility that venoms may 
have arresting effects beyond that of caus- 
ing temporary paralysis appears never to 
have been considered. 

Yu and Horstmann (1997) recognise 
eleven subgenera of Netelia of which four, 
Netelia s. str., Paropheltes Cameron, Besso- 
bates Townes, Townes and Gupta, and 
Parahates Foerster, are known to occur in 
Britain. This paper records the outcome of 
experiments to investigate possible long- 



Volume 10, Number 1, 2001 



103 



term venom effects in the first three of 
these subgenera of Netelia, and notes also 
other aspects of parasitoid behaviour and 
host development. 

The following species were used in ex- 
periments (comment on phenology and 
host range is supported by reared material 
in the National Museums of Scotland 
(NMS)): Netelia (Bessobates) cristata (Thom- 
son), a plurivoltine and normally solitary 
parasitoid of various exposed Noctuidae; 
N. (B.) laiuugula (Thomson), a univoltine, 
solitary and regular parasitoid of Opero- 
phtera hrumata (Linnaeus), sometimes 
reared also from other spring-feeding ar- 
boreal Geometridae; N. (Netelia) vimilae 
(Scopoli) (= cephalotes (Holmgren)), a uni- 
voltine, gregarious and regular parasitoid 
of Ceriira vinula (Linnaeus) (Notodonti- 
dae); N. (Paropheltes) tarsata (Brischke), a 
plurivoltine solitary parasitoid of sniall 
Geometridae, especially species of Eupithe- 
cia Curtis; and N. (P.) ?tliomsoni (Brauns), 
in Britain a widespread plurivoltine par- 
asitoid of Xanthorhoe fluctiiata (Linnaeus) 
and other similarly small to medium sized 
Geometridae. The latter comes closer to N. 
(P.) inedita (Kokujev) than to thovisoni in 
Delrio (1975), but Yu and Horstmann 
(1998) give the latter as a senior synonym. 
Delrio (1975) records thomsoiii from Geo- 
metridae including Eupithecia species. 

METHODS 

All livestock originated from England. 
Adult parasitoids were identified by ref- 
erence to Delrio (1975), and voucher spec- 
imens are deposited in the NMS. 

Netelia (Bessobates) cristata. One female 
(unmated) reared from Costuia trapezina 
(Linnaeus) from Hartslock, Oxfordshire, 
was used in experiments. The experimen- 
tal host, Lacanobia oleracea (Linnaeus), was 
reared from the previous generation in 
culture and fed on leaves of Taraxacum 
Weber and Crataegus Linnaeus. 

Netelia (Bessobates) latungula. Adults 
used in experiments were collected by 
sweeping Corylus Linnaeus at Hell Cop- 



pice, Buckinghamshire (two females), and 
reared from O. brumata from Gait Barrows 
NNR, Lancashire (one female, unmated). 
Experimental hosts were wild-collected 
larvae of O. brumata {Corylus, Hell Cop- 
pice; Corylus, Pamber Forest, Hampshire; 
and Salix Linnaeus, Otmoor, Oxfordshire), 
and cultured O. brumata and Theria pri- 
maria (Haworth) from the previous gen- 
eration. All hosts were fed on leaves of 
Crataegus. Some O. brumata larvae bearing 
eggs of N. latungula when collected were 
also investigated. 

Netelia (Netelia) vinulae. Parasitised lar- 
vae of C. vinula, collected at Druridge Bay, 
Northumbria, were received from H. A. 
Ellis (cf. ElHs 1998) and provided the adult 
N. vinulae used the following year. Five fe- 
males (at least two mated and at least two 
virgin) were involved in experiments. Lar- 
vae of the experimental host, C. vinula, 
were obtained from livestock dealers (ex 
culture) and fed on leaves of Salix. 

Netelia (Paropheltes) tarsata. One female 
(unmated) reared from Eupithecia absin- 
thiata (Clerck) from Sheffield, Yorkshire 
(received from T. H. Ford) was used in ex- 
periments. The experimental host, Eupithe- 
cia vulgata (Haworth), was cultured from 
the previous generation and fed on leaves 
of Crataegus. 

Netelia (Paropheltes) ?thomsoni. One fe- 
male (unmated) reared from Xanthorhoe 
fluctuata from Hampstead Heath, London 
(received from R. A. Softly) was used in 
experiments. Eupithecia natiata (Hiibner), 
cultured from the previous generation and 
fed on Calluna Salisbury, and Eupithecia 
sp. collected wild from Quercus Linnaeus 
at Pamber Forest, Hampshire were exper- 
imental hosts. 

All female parasitoids were seen to feed 
on honey: water (ca 1:3), with which they 
were kept at all times in corked glass 
tubes or clear plastic boxes (see below) 
apart from the brief periods of experimen- 
tal manipulation. Lepidoptera larvae were 
fed in ca 16 X 8 X 6cm closed clear plastic 
boxes lined with copious absorbent tissue 



104 



Journal of Hymenoptera Research 



paper. All livestock was kept in an un- 
heated, detached and fully shaded out- 
door shed under conditions of natural 
temperature and daylength except during 
experiments under observation. 

Experimental exposures of hosts to par- 
asitoids were done under observation (ex- 
cept for some with N. (B.) cristatn — see Re- 
sults) in 2.5 X 7.5 cm corked glass tubes 
(N. (B.) latungida and the two N. (Pnro- 
pheltes) species), or in ca 16 X 8 X 6 cm 
clear plastic boxes (N. (B.) cristatn and N. 
(N.) viiiulae). 

RESULTS 

Netelia (Bessobntes) cristatn. — Eggs were 
mature by the twelfth day after adult 
emergence. Egg dumping was not seen, 
probably because the single parasitoid ob- 
served was never deprived of hosts for 
long. This species is almost completely 
nocturnal and only four parasitisations 
(one with successful non-destructive con- 
current host-feeding — see Jervis and Kidd 
1986 for explanation of these terms) and 
one further non-destructive but non-con- 
current host-feeding event were observed 
directly. For oviposition the female 
grasped the late final instar host with all 
six legs, oriented head to tail, and moved 
backwards along the host's body towards 
its head, then rapidly oviposited on a tho- 
racic segment without separately stinging 
the host. In three cases the female then 
tried to bite the host mid-dorsally, pre- 
sumably in order to host-feed, but in each 
case the host writhed furiously and she 
was thrown off. In the only other ovipo- 
sition sequence observed the parasitoid 
laid a second egg, without releasing the 
host, before successfully host-feeding on 
haemolymph via a wound made with the 
mandibles. Host-feeding without oviposi- 
tion was observed on one host (earlier in 
its final instar) — as in oviposition sequenc- 
es the host was grasped tightly with all six 
legs, and the female chewed a wound at 
which she fed midway along the host's 
dorsum, at the same time curling the tip 



of her abdomen towards the host's thorac- 
ic segments but without ovipositing. 

Otherwise parasitised hosts were ob- 
tained by keeping the single female over- 
night with 5-7 hosts in a cardboard shoe 
box (ca 23 X 13 X 10 cm) closed with a 
sheet of glass, and removing the contents 
the following morning. Penultimate instar 
hosts, and those early in their final instars, 
never received eggs, but over a thirteen 
day period the female laid 54 eggs (50 of 
them in the last nine days) on late final 
instar larvae until the experiment was cur- 
tailed for want of further host larvae, al- 
though the female parasitoid lived for 
about another 55 days (ca 80 as an adult 
in all). At the end of a night, sixteen hosts 
had received one egg; nine had received 
two; two had received four; and one had 
received twelve eggs (generally, some 
hosts had also received no eggs, but as the 
smaller hosts were clearly less attractive 
there is no scope to analyse egg distribu- 
tion). The host with twelve eggs was 
reared and five parasitoid cocoons result- 
ed; the two hosts with four eggs similarly 
each resulted in two cocoons; two of four 
hosts reared with two eggs yielded two 
cocoons and the other two a single cocoon 
each; and all seven hosts reared with sin- 
gle eggs duly yielded the single parasitoid 
cocoon expected. This suggests that com- 
petition causes some mortality but that 
broods of two will regularly be fully via- 
ble. 

Nine hosts from which eggs were man- 
ually removed, and also one host bearing 
an egg that failed to hatch, produced pu- 
pae and then adult moths, apparently nor- 
mally. Two eggs on one host were allowed 
to hatch (which occurred two and five 
days after oviposition) and on the sixth 
day after oviposition the two larvae (four 
and one days old) were destroyed, after 
which the host pupated and became an 
adult moth. Two other hosts each bearing 
one parasitoid were reared for eight days 
after oviposition, by which time parasitoid 
larvae were ca six days old and ca three 



Volume 10, Number 1, 2001 



105 



times as long as the egg, before the para- 
sitoid larvae were killed. Both hosts be- 
came prepupal within two days, but died 
without pupating, apparently in an arrest- 
ed state. 

In all wild and experimental rearings 
permitted to reach such a state, the host 
was consumed as a prepupa in its pre- 
pared pupation site, in which the parasit- 
oid cocoon is constructed. The generation 
that overwinters does so in the cocoon. 

Netelia (Bessobates) latungula. — Eggs 
were mature five days after adult emer- 
gence, and started to be dumped ca ten 
days after emergence in the absence of 
hosts. If kept humid dumped eggs later 
split to reveal a living first instar larva, 
just as did eggs laid on hosts. Penultimate 
instar hosts (both O. hrumnta and T. pri- 
mnria) were rejected consistently, although 
sometimes investigated. Oviposition at- 
tacks on final instar hosts are evidently 
provoked by movement: hosts remaining 
still when contacted by the female were 
ignored. There was no pre-oviposition 
sting, the adult parasitoid rapidly pounc- 
ing on the host and ovipositing singly on 
one of the host's thoracic segments (very 
rarely at the host's caudal end: the two 
eggs laid in this position were quickly lost, 
possibly because the host could reach 
them with its mouthparts). After all seven 
ovipositions witnessed the host was then 
released, without the parasitoid attempt- 
ing to host-feed. Some interactions, how- 
ever, took a different course: in five cases 
the female parasitoid grasped a final instar 
O. bnimatn larva, jabbing it once with its 
ovipositor midway along its dorsum. Four 
of these larvae were then released and 
abandoned, but the parasitoid chewed a 
wound on the other from which it fed on 
haemolymph. No paralysis was evident. 

Eggs were cut off eleven parasitised O. 
briwmta larvae and two T. primaria, all of 
which pupated and produced adult 
moths. The five O. bruniata which had 
been possibly stung but not oviposited 



onto also pupated and produced adult 
moths apparently normally. 

In a heavily parasitised field sample of 
final instar O. brumata larvae collected 
from Con/lus on 7.vi.l979 at Hell Coppice, 
Buckinghamshire, 15 bore no eggs, 19 had 
one egg, and nine carried two eggs of N. 
(B.) Ititufigula. Five of the latter nine pro- 
duced single N. (B.) latungula cocoons (the 
other four produced other parasitoids pre- 
emptively), showing that this species is 
probably strictly solitary but apparently 
incapable of rejecting already parasitised 
hosts. 

In all wild and experimental rearings 
permitted to reach such a state, the host 
was consumed as a prepupa in its pre- 
pared pupation site, and the parasitoid 
overwintered there in its own cocoon. 

Netelia (Netelia) viuulae. — Two copula- 
tions each of about four minutes duration 
were observed: in both cases the male 
climbed on the dorsum of the female, 
which was standing on a horizontal sur- 
face, so that the copula was orientated 
head to head. This contrasts with the ac- 
count of copulation given by Vance (1927) 
for N. (N.) spinipes (Cushman). 

Female parasitoids contained mature 
eggs from about the eighth day after their 
emergence, and by the eleventh day fe- 
males having had no access to hosts start- 
ed to dump eggs on the sides of the con- 
tainer. Such eggs, if kept humid, later split 
to reveal the first instar larva, just as eggs 
placed on hosts. The females were offered 
both penultimate and final instar larvae, 
attacking the former (including those in 
proecdysis/apolysis between these in- 
stars) very much more readily. They ad- 
vanced on hosts at first rather cautiously, 
making repeated stinging attacks with 
brief (rarely as long as a second) insertions 
of the ovipositor, usually in central posi- 
tions along the host's body. The hosts, 
which at first attempted to defend them- 
selves by thrashing and sometimes daub- 
ing oral secretions, were gradually sub- 
dued; those in the penultimate instar in 



106 



Journal of Hymenoptera Research 



particular becoming markedly incapacitat- 
ed, though not fully paralysed. The female 
then sometimes rejected hosts (in both 
penultimate and final instars) in which she 
had invested considerably (as many as ten 
stinging attacks) without ovipositing, or 
else she climbed onto the host and com- 
menced oviposition onto thoracic seg- 
ments. Each site for anchoring an egg ap- 
peared to be chosen with some delibera- 
tion. After the ovipositor tip was inserted 
through the host's cuticle it took approxi- 
mately half a second for the egg to slide 
down the outside of the ovipositor to be 
anchored into the epidermis by its stalk, 
which travels down the ovipositor shaft 
internally. Host movement as it recovered 
during the egg-laying process greatly in- 
hibited the female, and this may impose a 
strong limit to brood size (about 4-8 eggs 
seems usual). Following oviposition, the 
female usually fed on host haemolymph 
through a wound which it chewed with 
its mandibles mid-way along the host's 
dorsum. Hosts that were stung but then 
rejected were not fed on in this way, and 
haemolymph oozing from stinging sites 
was not imbibed. The behaviour of fe- 
males towards hosts before their penulti- 
mate instar was not investigated. During 
host ecdysis eggs anchored into the epi- 
dermis of penultimate instar hosts easily 
tear through the cuticle being sloughed 
and losses are normally extremely low. 

Three penultimate instar hosts that had 
suffered stinging attacks without oviposi- 
tion, and three penultimate instar hosts on 
which oviposition had also occurred but 
the progeny was destroyed, were reared 
to investigate the effects of venom. All six 
pupated and produced adult moths con- 
temporaneously with controls. 

In all wild and experimental rearings 
permitted to reach such a state, the host 
was consumed as a prepupa in its pre- 
pared pupation site, and the parasitoids 
overwinter there in their own cocoons. 

Netelia (Paropheltes) tarsata. — The female 
parasitoid lived as an adult from 5.vi until 



24. ix, a period of HI days. Hosts were not 
offered until the parasitoid's 60th day of 
adult life. An unknown number of eggs 
had been dumped but, once experiments 
commenced, further egg dumping was not 
seen, probably because the female was not 
then deprived of hosts. Indeed, on one oc- 
casion an egg that was clearly immature 
(greenish grey rather than the usual black 
in colour) was laid on a host, but this and 
also several others of the eggs laid failed 
to anchor adequately and were lost, pre- 
sumably because the chorion had not suf- 
ficiently hardened. The eggs are more or 
less coriacious and matt, unlike the shiny 
eggs seen in the species of the subgenera 
Netelia and Bessobates here studied. Final 
instar hosts were greatly preferred, 
though some ovipositions occurred on 
penultimate instar hosts (including one in 
the proecdysis/apolysis preceding the fi- 
nal instar). Hosts were stung once, with an 
insertion of the ovipositor into an abdom- 
inal position, causing partial temporary 
paralysis. The parasitoid then oviposited 
once onto a thoracic segment, and subse- 
quently nearly always chewed a wound 
midway along the host's back from which 
it fed on haemolymph. In one case host- 
feeding commenced before oviposition 
and continued during it and for a time 
afterwards. Host-feeding was often so 
heavy (green fluid greatly distending the 
parasitoid's metasoma) that the female 
was unable to move with agility and did 
not attack further hosts until the following 
day. A limited capacity to deal with more 
than two hosts per day was also evi- 
denced by her laying immature eggs (see 
above) and sometimes failing to produce 
enough venom to subdue hosts to the 
point that she was willing to oviposit on 
them. 

Three penultimate instar and two final 
instar hosts that had received paralysing 
venom but no eggs, and three further final 
instar hosts that had also received eggs 
which were then removed, all pupated 
and produced adult moths apparently 



Volume 10, Number 1, 2001 



107 



normally. An additional four final instar 
and one penultimate instar hosts from 
which eggs had been removed died of a 
fungal disease that also affected some un- 
exposed control larvae. 

In all wild and experimental rearings 
permitted to reach such a state, the host 
was consumed as a prepupa in its pre- 
pared pupation site, and the generation 
that overwinters does so in the cocoon. 

Netelia (Pnropheltes) Ithomsoni. — Eggs 
(which are similar to those of N. (P.) tar- 
satn) were dumped after several days of 
host deprivation. Two similar ovipositions 
were observed, on a final instar Eupithecia 
tmnatn and on a final instar Eupithecia sp. 
The female stung the host (10-20 seconds 
insertion) in an abdominal position caus- 
ing rather full temporary paralysis; she 
then oviposited singly onto a thoracic seg- 
ment, when the body of the egg was seen 
to slide down the outside of the ovipositor 
during ca 1 second; and subsequently she 
chewed a wound midway along the host's 
back from which she fed on haemolymph. 
The cultured host (£. nanata) died of a fun- 
gal disease and the other (wild collected) 
host proved to be already parasitised. No 
further trials could be conducted. 

In all wild rearings the host was con- 
sumed as a prepupa in its prepared pu- 
pation site, and the generation that over- 
winters does so in the cocoon. 

DISCUSSION 

Pre-ovipostion venoms causing a de- 
gree of temporary paralysis have been re- 
ported in several species of the subgenus 
Netelia previously (She vy rev 1912, Cush- 
man 1926, Vance 1927) and studies on spe- 
cies of the subgenus Bessobates have either 
stated that no paralysis is caused (Shevy- 
rev 1912) or been reported without men- 
tion of it (Stenton 1910, whose account of 
his parasitoid of O. brumata almost cer- 
tainly refers not to N. (B.) virgata but to N. 
(B.) latiDigula, as it is clear from material 
in NMS that C. Morley, who had deter- 
mined Stenton's specimens, regularly mis- 



identified latungula as virgatus). The pres- 
ent study concurs with these earlier find- 
ings, but the biology of the subgenus Par- 
opJieltes does not seem to have been 
previously studied even to this extent. 

The main finding of the present work is 
that no substances controlling host devel- 
opment are injected during the oviposi- 
tion process in any of the three subgenera 
investigated, since hosts from which eggs 
had been removed developed normally. 
Thus the venoms injected prior to ovipos- 
iton in the subgenera Netelia and Parophel- 
tes have no effect on the host other than 
to subdue it, simply allowing the female 
parasitoid to place eggs accurately using 
its appreciably exserted ovipositor. Spe- 
cies of the subgenus Bessobates, which 
have markedly shorter ovipositors, ovi- 
posit more rapidly without immobilising 
the host other than physically by grasping 
it. Oviposition has been described for one 
species in the genus Phytodietus, the pu- 
tative sister genus to Netelia, and a venom 
causing temporary paralysis, often requir- 
ing several insertions of the ovipositor as 
in N. (N.) vifmlae, was observed (Sim- 
monds, 1947). This may suggest that the 
use of a paralysing venom is plesiomorph- 
ic in Netelia (see also Kasparyan 1973 
[1981:49]). 

In all tribes of Tryphoninae other than 
Idiogrammatini and Phytodietini eggs 
ready for oviposition are commonly car- 
ried externally on the ovipositor until a 
host is found (pers. obs.; but for Ankylo- 
phonini and Sphinctini, A. Bennett pers. 
comm.). In the studied subgenera of Ne- 
telia (Phytodietini) this behaviour is not 
approached: in all three subgenera the egg 
did not start to issue from the parasitoid's 
genital opening until the ovipositor was 
inserted for attaching the egg to the host. 
Although observations of oviposition in 
the tribes carrying eggs on the ovipositor 
are rather sparse, only the exenterine Ex- 
etiterus abruptorius (Thunberg) has been re- 
ported to cause temporary paralysis (Mor- 
ris 1937). This observation would indicate 



108 



Journal of Hymenoptera Research 



that carrying the egg on the ovipositor 
(which it is here presumed to be the case 
in E. abruptorius, though it is not expHcitly 
stated to be so by Morris) does not pre- 
clude the use of a paralysing venom, but 
the lack of records of paralysis by Exen- 
terini and Tryphonini stemming from oth- 
er studies (see Kasparyan 1973 [1981:49]) 
suggests that it is not usual in these tribes. 
The situation is uncertain in Idiogram- 
matini: despite some speculation by Cush- 
man (1937) and an erroneous citation by 
Kasparyan (1973 [1981: 49]), the process of 
oviposition in Idiogramma Foerster seems 
not to have been reported, but eggs have 
not been found to be carried externally on 
the ovipositor (A. Bennett pers. comm.). 

Thus there remains no evidence that 
Tryphoninae deploy venom systems that 
have effects other than to subdue the host 
while oviposition takes place, although 
egg-removal experiments have been con- 
ducted only for Netelia and should be car- 
ried out for other tribes. Furthermore the 
other two known groups of koinobiont ec- 
toparasitic Ichneumonidae, i.e. some Ade- 
lognathinae and the Poh/sphincta genus- 
group, appear to employ pre-oviposition 
venoms that, as in two of the subgenera 
of Netelia studied here, have only this sub- 
duing effect. Adelognathinae consists of 
the single genus Adelognnthus Holmgren. 
One species, A. chn/sopi/gns (Gravenhorst) 
(= grmuilatus Perkins), is a typical idio- 
biont, employing venom for the perma- 
nent paralysis of its host (Rahoo and Luff 
1987, confirmed by F.D. Bennett pers. 
comm.). However, most Adelognnthus spe- 
cies appear to be koinobionts (Fitton et al. 
1982), and one thoroughly investigated 
but undescribed species employs a venom 
causing temporary paralysis, facilitating 
oviposition, but which has no other ap- 
parent effect: hosts intercepted after pa- 
ralysis but before oviposition, as also hosts 
from which eggs were removed, recover 
to develop to the adult stage normally 
(Shaw unpublished). In the Polysphincta 
genus-group temporary paralysis of the 



host prior to oviposition is probably com- 
mon to all genera and has been described 
by Cushman (1926) for Zatypota parva 
(Cresson) and by Eberhard (2000a) for Hy- 
menoepicnemis argyraphaga Gauld. In the 
latter species Eberhard (2000a) found that 
the venom appears to have no long tern^ 
effect on host development. This appears 
to be generally true of the Polysphincta ge- 
nus-group: I have on several occasions 
reared to adulthood immature spiders 
bearing parasitoid larvae at the time of 
collection that for one reason or another 
subsequently died young, to check that 
host development was unimpaired, and in 
every case (involving the parasitoid gen- 
era Polysphincta, Dreisbachia Townes, Sch.i- 
zopyga Gravenhorst, Zatypota Foerster and 
Acrodnctyla Haliday) it was found to be so. 
Thus, although more investigation of Try- 
phoninae is warranted, it appears that 
none of the extant groups of koinobiont 
ectoparasitic Ichneumonidae employs a 
venom system with an effect remotely 
similar to that seen in the cyclostome Bra- 
conidae, which seems to have been so im- 
portant in the evolution of endoparasitism 
within at least one lineage (Shaw 1983, 
Whitfield 1992). 

Whereas the two studied species of the 
subgenus Bessobates will normally oviposit 
only on final instar larvae, and one species 
clearly avoided inactive hosts, the single 
species studied in the subgenus Netelia, N. 
(N.) viniilae, which is unusual in both the 
subgenus and the genus as a whole for be- 
ing fully gregarious, preferred hosts in the 
penultimate instar (see also Ellis 1998) and 
was very willing to attack them when they 
were inactive in the proecdysis/apolysis 
between that and the final instar. Other 
studied species of the subgenus Netelia are 
solitary, and, although apparently prefer- 
ring final instar hosts, also commonly ovi- 
posit on penultimate instar hosts (Guppy 
1961, Danks et al. 1979). At least one of the 
species of the subgenus Paropheltes inves- 
tigated here also seems to be somewhat 
plastic in this respect. 



Volume 10, Number I, 2001 



109 



Species of all three subgenera dumped 
eggs in the absence of hosts, as has been 
observed before in the subgenera Netelia 
(Vance 1927) and Bessobntes (Stenton 1910). 
This presumably happens because embry- 
onic development is independent of the 
host, contrary to Stenton's (1910) view, 
and eggs in these two subgenera (not in- 
vestigated for Pnropheltes) can "hatch" 
even if dumped. The degree of maturity 
of eggs that are laid on hosts varies con- 
siderably and this may have a bearing on 
the time that elapses before they hatch, al- 
though, as dumped eggs tend to dry up 
without hatching unless kept humid, pre- 
vailing humidity is clearly also important 
and may explain why, in the wild, eggs of 
many Tryphoninae typically hatch only 
after the host has constructed its pupation 
chamber (cf Clausen, 1932). An egg-hatch- 
ing/larval development response to the 
conditions of high humidity developing in 
the host cocoon thus may be the principle 
means of ensuring the commencement of 
parasitoid development at the appropriate 
host stage in Tryphoninae, in contrast to 
koinobiont ectoparasitoid taxa such as 
Rhi/sipolis (Shaw 1983) and the eulophid 
Euloplius ramicornis (Fabricius) (Shaw 
1981, as E. Inrvarum (Linnaeus)) which 
achieve this through their venoms. Hosts 
parasitised by Netelia collected while they 
are still feeding in the wild almost invari- 
ably bear only unhatched eggs (pers. obs.), 
though once brought under captive con- 
ditions involving the high humidity of 
closed containers the eggs frequently 
hatch, and the larvae start to develop, be- 
fore the host constructs its pupation cham- 
ber. In extreme cases (cf. that illustrated 
by Ellis 1998 for N. (N.) vimilae) this can 
lead to the host being overwhelmed pre- 
maturely and a failure of the parasitoids 
to survive. The extent to which larval de- 
velopment per se may depend on high hu- 
midity would be worth investigating. 

Guppy (1961) and, possibly following 
him, Danks et al. (1979), writing on species 
in the subgenus Netelia, have stated that 



the host's development is "arrested" as a 
prepupa as the parasitoid starts to feed, 
but without giving reasons for that con- 
clusion. In one case for one species of the 
subgenus Bessobates studied, two parasit- 
oid larvae on one host were allowed to 
feed for up to 4 days before they were 
killed, and the host then went on to pu- 
pate normally. In two further cases in- 
volving this host and parasitoid species 
the parasitoid larvae were killed after 6 
days, when they were still too small to 
have caused much damage to the larval 
hosts, and both hosts then burrowed for 
pupation but died as prepupae in what 
seemed to be an arrested state. The inter- 
esting possibility that the larva of Netelia 
species does indeed inhibit host develop- 
ment, other than by causing damage 
merely in the course of feeding, is worthy 
of further investigation, though it appears 
not to be capable of this in the earliest 
days of its feeding. Eberhard (2000a, b) 
gives evidence of control over the host, 
apparently through chemicals produced 
by the parasitoid larva, in Hymenoepimecis 
argyraphaga of the Polysphincta genus- 
group. 

The use of mandibles, rather than the 
ovipositor, to make the wound necessary 
for non-destructive host-feeding has been 
noted before in the subgenus Netelia 
(Vance 1927), and was common to all 
three of the subgenera studied. In one sub- 
genus, Bessobates, it appears sometimes (or 
possibly usually) to involve host individ- 
uals other than those used for oviposition, 
perhaps because a different grip on an un- 
subdued host is required. In contrast, in 
the subgenera Netelia and Paropheltes non- 
destructive host-feeding is concurrent (cf. 
Jervis and Kidd 1986) and normally occurs 
following each oviposition. The behaviour 
in Netelia s.l. is in marked contrast to that 
recorded by Simmonds (1947) for Phyto- 
dietus nifipes pulcherrimus (Cresson), in 
which feeding was said to take place on 
the fluids that exuded from the puncture 
wound (or wounds) made by the ovipos- 



110 



Journal of Hymenoptera Research 



itor in the course of injecting venom, and 
furthermore if the adult fed from the host 
it did so before ovipositing on it, rather 
than afterwards as in the subgenera Nete- 
lia and (normally) Pnroplieltes. Remark- 
ably, Zinnert (1969) noted that Erromenus 
calcator (Miiller) (Tryphonini) wounds the 
host with its mandibles but does not then 
feed on the exuding haemolymph. Other 
Ichneumonidae known to make feeding 
wounds with their mandibles include the 
tryphonine Eridolius rufonotatiis (Holm- 
gren) (Exenterinae) (Carl 1976), the pimp- 
line Exeristes rohorntor (Fabricius) (Fox 
1927) and an undescribed species of Ade- 
lognatlius (Shaw unpublished). Elsewhere 
in the Hymenoptera it is known in Eulo- 
phidae, Scelionidae and particularly in 
parasitoid aculeate groups (cf. Jervis and 
Kidd 1986). 

ACKNOWLEDGMENTS 

I am grateful to Hewett Ellis, Tom Ford and Ray 
Softly for giving me parasitoid livestock, to Rob Pe- 
tley-Jones and English Nature for permission to do 
fieldwork at Gait Barrows NNR, to the Berkshire, 
Buckinghamshire and Oxfordshire Naturalist's Trust 
for allowing my fieldwork on their nature reserves, 
and to Andy Bennett and David Wahl for construc- 
tive reviews of a previous draft. 

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X 
^aClETy 




Journal of 

Hymenoptera 
Research 



Volume 10, Number 2 October 2001 

ISSN #1070-9428 
CONTENTS 

CAETANO, F. H. and F. J. ZARA. Ultramorphology and histology of the foregut and 
midgut of Pachycondyla (= Neoponera) villosn (Fabricius) larvae (Formicidae: 
Ponerinae) 251 

GOOLSBY, J. A., C. J. BURWELL, J. MAKINSON, and R DRIVER. Investigation of the 
biology of Hymenoptera associated with Fergusonina sp. (Diptera: Fergusonini- 
dae), a gall fly of Melaleuca quinquenervia, integrating molecular techniques 163 

GRISSELL, E. E. and G. L. PRINSLOO. Seed-feeding species of Megastigmus (Hymenoptera: 

Torymidae) associated with Anacardiaceae 271 

ODA, R. A. M., M. V. de MAC EDO, and D. L. J. QUICKE. First biological data for 
Aspilodeiiion Fischer (Hymenoptera: Braconidae: Hydrangeocolinae): parasitoids of 
cecidomyiid fly galls on Asteraceae in Brazil 126 

OHL, M. The southern African wasp genus Handlirschia Kohl, 1897 (Hymenoptera: 

Apoidea, Sphecidae, Bembicinae) 231 

PRINSLOO, G. L. The Afrotropical species of Leptomastidea Mercet (Hymenoptera: 

Encyrtidae), parasitoids of mealybugs 145 

QUICKE, D. L. J. and A. PURVIS. A new species of Streblocera {Asiastreblocera) 

(Braconidae: Euphorinae) from Thailand with depressed ovipositor 138 

SCHAUFF, M. E. and D. H. JANZEN. Taxonomy and ecology of Costa Rican Euplectrus 

(Hymenoptera: Eulophidae), parasitoids of caterpillars (Lepidoptera) 181 

SCHIFF, N., A. J. FLEMMING, and D. L. j. QUICKE. Spermatodesmata of the sawflies 

(Hymenoptera: Symphyta): evidence for multiple increases in sperm bundle size ... 119 

, JUN 2 7 2002 

(Continued on back cover) 



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J. HYM. RES. 
Vol. 10(2), 2001, pp. 113-118 

A New Species of the Genus Orussonia Riek and the Female of 
O. depressa Riek (Hymenoptera: Symphyta, Orussidae) 

Stefan Schmidt and Gar^ A. P. Gibson 

(SS) Australian National Insect Collection, CSIRO Entomology, Canberra ACT 2601, Australia 

(e-mail: Stefan.Schmidt@ento.csiro.au); (GAPG) Systematic Entomology Section, Eastern Cereal 

& Oilseed Research Centre, Agriculture and Agri-Food' Canada, K.W. Neatby Building, 960 

Carling Avenue, Ottawa, Ontario, Canada, KIA 0C6 (e-mail: gibsong@em.agr.ca) 



Abstract. — Orussonia rnficaiidata Schmidt and Gibson, new species, and the female of O. de- 
pressa Riek are newly described; the male of O. depvessa is redescribed and differential features of 
the species and of the sexes of O. depressa are illustrated. 



Riek (1955) established the genus Onis- 
souia for two males collected in 1952, which 
he described as O. depressa. One of us 
(GAPG) recently collected two females that 
we associate as the opposite sex of O. de- 
pressa. We also discovered rri the Austra- 
lian National Insect Collection a third fe- 
male, collected many years earlier in 1919, 
which Riek had identified as a second un- 
described species and labelled with the 
manuscript name O. rnficaiidata. Here we 
describe what we consider to be the female 
of O. depressa and the female of this second 
species, which we name O. rnficaiidata. 

Orussonia is one of three genera of Orus- 
sidae occurring in Australia (Riek 1955). 
Tlie other two genera are Giii^lia Benson 
with at least four Australian species (Ben- 
son 1955, Riek 1955, Vilhelmsen, pers. 
comm.) and Ornssobains with six species 
from Australia and one from New Guinea 
(Schmidt and Vilhelmsen, in prep.). The 
genus Orussonia constitutes, together with 
genera traditionally placed in the tribe Lep- 
torussini by Benson (1955), a paraphyletic 
group comprising the basalmost lineages 
of the Orussidae (Vilhelmsen, pers. 
comm.). Females of species of the genus 
Orussonia possess two pairs of medially 
separate coronal teeth (Figs 1, 3) and an in- 
complete ventral transverse frontal carina. 



whereas in most other orussid genera there 
are at least three pairs of medially separate 
coronal teeth and a fully developed ventral 
transverse frontal carina (Vilhelmsen, pers. 
comm.). Among all orussids Orussonia is 
the only genus with an extremely dorso- 
ventrally flattened body and a prognathous 
head and is, therefore, easily recognisable. 
Nothing is known about the biology of 
Orussonia, but the flattened body shape 
might indicate that they crawl under loose 
bark. It is known for some beetles with 
dorsoventrally flattened bodies that they 
occur under bark, e.g. species of the genus 
Platisus (Coleoptera, Cucujidae) 
(Lawrence and Britton 1991). 

METHODS 

The holotype female and male of O. riif- 
icaudata and O. depressa, respectively, were 
used for the scanning electron micro- 
graphs. All specimens were uncoated and 
digital images were obtained with a Phil- 
ips XL 30 ESEM. The low vacuum envi- 
ronmental scanning mode was used for all 
except Fig. 1, which was obtained under 
high vacuum. A 1 mm cone aperture was 
used during low vacuum mode to im- 
prove image quality, which resulted in a 
circular image format at low magnifica- 
tions. The digital images were enlianced 



114 



Journal of Hymenoptera Research 




Figs. 1-6. OniSbOiiia species. 1-2, O. nificaiidatn, sp. n.: I, he.id, frontolateial \ lew; 2, face, v-h, ( ». ilcpn'Sfiii 
Riek: 3, head of female, frontolateral view; 4, face of female; 5, head of male, frontal view; 6, face of male, 
frontolateral view. Scale bars = |xm. 



Volume 10, Number 2, 2001 



115 



and the final plates compiled using Adobe 
Photoshopi^. Figure 10 is a composite of 
two images and figure 11 is a composite 
of four images. 

Orussonia Riek 

Orussonia Riek 1955: 104. 

Type species: Orussonia depressa Riek, by origi- 
nal designation. 

Description. — Body strongly dorsoven- 
trally depressed, thorax height about half 
its maximal width measured from above. 
Head prognathus, occiput strongly exca- 
vated and appressed over convex anterior 
margin of pronotuni. Vertex of female 
with two, of male with three separate 
pairs of coronal teeth and a less conspic- 
uous, smaller, subcontiguous medial pair 
dorsally of the lateral ocelli. Female anten- 
na 10-segmented; eighth and ninth seg- 
ments separated by a fine suture and 
forming a club; tenth segment peg-like, 
apically truncate and arising near middle 
of inner margin of ninth segment. Male 
antemia filiform, 11-segmented. Antenna 
about 1.5 X as long as head width, third 
segment shorter (0.56-0.71) than com- 
bined length of fourth and fifth segments, 
sixth segment slightly longer than third. 
Face between compound eyes flat with 
shallow punctures, without longitudinal 
carinae (Figs. 1-6). Gena rugose. Episto- 



mal sulcus absent, ventral transverse fron- 
tal carina reflexed and with shallow me- 
dian incision. Maxillary palp 5-segmented, 
labial palp 3-segmented. Mesoscutum and 
scutellum flat, forming a plane; scutellum 
crenulate along scutoscutellar line but oth- 
erwise smooth and shiny except for a few 
small setiferous punctures (Figs. 7, 8). Ax- 
illae punctate-rugose, contiguous medial- 
ly, separated by a distinct median carina 
(Figs. 7, 8). Mesopleuron laterally reticu- 
late-rugulose to punctate and setose ex- 
cept for smc^oth and shiny bare region me- 
dially below base of hind wing; ventrally 
much smoother with scattered setiferous 
punctures except more coarsely sculp- 
tured anterolaterally (Fig. 11). Metafemur 
swollen, about 2.5 x as long as its maxi- 
mal width. Forewing hyaline behind stig- 
ma but partly infuscate basally and api- 
cally, with two cubital cells, without inter- 
costal crossvein; anal cell closed, petiolate. 
Radial cell of forewing and hind wing 
open apically. Hind wing with large jugal 
lobe and without closed medial cell. Ab- 
dominal tergites 2-5 and 6 basally, later- 
ally carinate; second tergite subequal in 
length or slightly shorter than combined 
length of third and fourth tergites. 

Reumrks. — Orussonia is readily distin- 
guished from all other orussid genera by 
its strongly depressed body and progna- 
thus head (Riek 1955, flgs. 2, 3). 



KEY TO SPECIES OF ORUSSONIA RIEK 

1 Male and female: face with punctures separated by distinct interspaces, some of which are 
greater than the diameter of a puncture (Figs. 3-6); abdomen black, apical segment occasionally 
orange-brown depressa Riek 

- Female: face with punctures nearly contiguous, separated by narrow interspaces (Figs. 1, 2); 
abdomen black with apical two segments orange-brown ruficaudata, new species 



Orussonia depressa Riek 

(Figs. 3-11) 

Orussonin depressa Riek 1955: 104-5. Holotype 
6, Australia, NSW, 4 miles N. Bateman's 
Bay, 14.X.1952, E.F. Riek. Type in the Austra- 



lian National Insect Collection, Canberra. 
Paratype S, same data as holotype, in The 
Natural History Museum, London (both ex- 
amined). 

Additional materml examined. 2 9 9, Austra- 



116 



Journal of HvMHNorxERA Research 




Figs. 7-11. OniSMiiin dcprcssn Rick. 7, femak" thor-dv, dorsal view; <S, male thorax, dorsal \ievv; '■>, abdominal 
tergites 1 and 2 of female, dorsal view; 1(1, abdominal tergites 1 and 2 of male, dorsal \iew; 11, thorax and 
base of abdomen of male, ventrolateral view. Scale bars = (xm. 



Volume 10, Number 2, 2001 



117 



lia, ACT, Canberra, Black Mountain, 
2.xi.l998 and 23.X.1998, G.A.P. Gibson. 
The specimens were collected on dead eu- 
calypt trees. One specimen deposited in 
ANIC and one in the Canadian National 
Insect Collection, Ottawa. 1 9, Victoria, 
Red Hill, Oct. 1965, D.R. Holmes, in Ob- 
erosterreichisches Landesmuseum, Linz, 
Austria. 

Female. — Body length 5-6 mm. Black 
with following yelk^wish to orange- 
brown: labrum and labiomaxillary com- 
plex, tenth flagellar segment and some- 
times basal three to seven flagellar seg- 
ments, protibia apically on outer surface, 
tarsi except apical segment sometimes 
brown, metatibia dorsomedially, abdomi- 
nal sternites medially and apical tergite 
occasionally. Forewing smoky-hyaline 
with band near base and larger region be- 
yond stigma brownish; stigma black. Face 
(Figs. 3, 4) flat with shallow, oval to sub- 
circular punctures separated by smooth, 
shiny, glabrous interspaces, the largest in- 
terspaces greater than diameter of a punc- 
ture; punctures each with single seta. Me- 
soscutum (Fig. 7) punctate-rugose except 
with shiny, smooth or mostly sniooth me- 
dial band, and with a less ciistinct smooth 
region laterally near parapsidal line. Ab- 
domen with basal two tergites reticulate- 
rugose except medially and apically 
smooth to finely coriaceous (Fig. 9); re- 
maining tergites primarily coriaceous but 
with increasingly obscure, paramedial, 
transverse punctate-rugose line anteriorly 
on tergites 3-5. 

Male. — Similar to female except for fol- 
lowing: face (Figs. 5, 6) densely setose me- 
dially over region about as wide as clyp- 
eus (Fig. 5), flat interspaces with numer- 
ous setae originating from distinct setal 
pores (Figs. 5, 6); mesoscutum (Fig. 8) 
punctate-rugose except punctate with 
smooth interspaces anteriorly and with 
obscure median carina. Second abdominal 
tergite without medial coriaceous band 
and with larger and more distinct shiny 
spot basolaterally (Fig. 10). 



Remarks. — We consider it likely that the 
described differences between the sexes 
are secondary sexual features. This would 
be consistent with sexual differences 
found in other orussids. However, we are 
currently unable to exclude the possibility 
that the females and males represent sep- 
arate species. Discovery of the male of O. 
ruficaudata would provide more informa- 
tion on sexual differences among species 
of Onissouia. 

Onissonia ruficaudata Schmidt and 
Gibson, new species 

(Figs. 1, 2) 

Holotype female: [Australia] "Lakes En- 
trance, October 1919, V. [Victoria], F.E. 
Wilson"; "Ex. Coll. Nat. Mus"; "Holotype 
Orussoiiia ruficaudata Riek"; "manuscript 
name"; "Holotype Orussoiiia ruficaudata 
Schmidt & Gibson". Condition of holo- 
type: right antenna missing beyond scape; 
mounting pin through mesoscutum me- 
dially. Type deposited in the Australian 
National Insect Collection, Canberra. 

Female. — Body length 8.5 mm. Dark 
brown to black with following yellowish 
to orange-brown: labrum and labiomaxil- 
lary complex, antenna except segment 
nine and apical margin of segment eight, 
tibiae and tarsi except meso- and metati- 
bia darker basally, apical two abdominal 
segments entirely and sternites 3-5 large- 
ly. Forewing hyaline with brownish re- 
gion behind costal vein and large brown- 
ish region beyond stigma; stigma black. 
Face (Figs. 1, 2) flat with shallow, dense, 
almost alveolate punctures separated by 
interspaces which are narrower than di- 
ameter of a puncture; punctures each with 
single seta. Vertex with distinct, dark red 
coronal teeth. Mesoscutum punctate, 
punctures of different sizes and partly 
confluent, and with smooth band medially 
and paralaterally near parapsidal line. Ab- 
domen with basal two tergites rugulose 
except medially and apically coriaceous to 
imbricate; remaining tergites primarily re- 
ticulate-coriaceous but with obscure, par- 



118 



Journal of Hymenoptera Research 



amedial, transverse punctate-rugose line 
anteriorly on tergites 3-5. 

Male. — Unknown. 

Remarks. — In addition to the two fea- 
tures used to differentiate females of O. 
riificaudata and O. depressa in the key, the 
single female of O. riificaudata also has 
more distinctly differentiated punctures 
on the mesoscutum. The two O. depressa 
females have a more rugose mesoscutum. 

ACKNOWLEDGEMENTS 

We thank Ms Jennifer Read (Eastern Cereal & Oil- 
seed Research Centre) who produced the scanning 
electron photomicrographs and the plates of illustra- 
tions. We also would like to thank Dr. Lars Vilhelm- 



sen (Department of Systematic Zoology, Uppsala, 
Sweden) for discussing orussid relationships. A spec- 
imen for this study was provided by Dr. Fritz Gus- 
enleitner (Oberosterreichisches Landesmuseum, Linz, 
Austria). 

LITERATURE CITED 

Benson, R. B. 1955. Classification of the Orussidae 
with some new genera and species (Hymenop- 
tera; Symphyta). Proceedings of the Roi/al Eiitoiiio- 
logical Societi/ of Loudon (B) 24, 13-23. 

Lawrence, J.F. and E.B. Britton. 1991. Coleoptera. In; 
CSIRO Entomology (Ed.). The Insects of Australia. 
Second Edition. Melbourne University Press, 
Melbourne, pp. 543-683. 

Riek, E.F. 1955. The Australian sawtlies of the family 
Orussidae (Hymenoptera, Symphyta). Australian 
jourjial of Zoologxj 3: 99-105. 



J. HYM. RES. 
Vol. 10(2), 2001, pp. 119-123 

Spermatodesmata of the Sawflies (Hymenoptera: Symphyta): 
Evidence for Multiple Increases in Sperm Bundle Size 

Nathan Schiff, Anthony J. Flemming, and Donald L. J. Quicke 

(NS) U.S. Forest Service, Southern Research Station, Center for Bottomland 

Hardwoods Research, P.O. Box 227, Stoneville, MS 38776, USA; (AJF) Department of Biology, 

hnperial College at Silwood Park, Ascot, Berkshire SL5 7PY, U.K.; (DLJQ) Unit of Parasitoid 

Systematics, Department of Biology, Imperial College at Silwood Park, Ascot, Berkshire SL5 

7PY, UK, and Department of Fntomology, The Natural History Museum, 

London SW7 5BD, UK 



Abstract. — We present the first survey of spermatodesmata (bundles of spermatozoa connected 
at the head by an extracellular 'gelatinous' matrix) across the sawfly superfamilies. Spermatodes- 
mata occur in all examined taxa within the sawfly grade (Xyelidae-Orussidae inclusive), but are 
not found in the Apocrita. Using DAPI staining, the numbers of indivicUial sperm per sperma- 
todesm were calculated and the values obtained are mapped on to the current phylogenetic hy- 
pothesis. The plesiomorphic spermatodesm in the Hymenoptera, based on that observed in the 
putatively basal family Xyelidae, contains relatively few sperm, approximately 16. However, in 
the Tenthredinoidea and in the Siricidae, far larger numbers are found, reaching up to 256 in the 
Cimbicidae. 



In many insects, mature sperm released 
from testicular follicles are neither free in- 
dividuals nor packaged into variously 
complex spermatophores, but are ar- 
ranged in organised bundles with their 
anterior ends embedded in an extracellu- 
lar cap. These structures, called sperma- 
todesmata (spermatodesm singular), oc- 
cur, amongst others, in at least some men"i- 
bers of the Collembola, Orthoptera, Dip- 
tera, Coleoptera, Lepidoptera and 
Hymenoptera (Jamieson 1987). Within the 
Hymenoptera, spermatodesmata appear 
to be limited to the basal sawflies (Quicke 
et al. 1992, Quicke 1997), and they have 
not been observed in members of the 'Eva- 
niomorpha' (Stephanoidea, Megalyroidea, 
Evanioidea and Ceraphronoidea exam- 
ined), proctotrupoid sensii lata (Diapriidae, 
Proctotrupidae, Heloridae and Scelionidae 
examined), chalcidoid, cynipoid, ichneu- 
monoid or aculeate groups (Quicke et al. 
1992, Newman and Quicke 1998, 1999a,b, 
2000, Lino-Neto et al. 1999, 2000a,b). 



Until now, spermatodesmata have only 
been characterised in a few sawflies, al- 
most entirely as part of ultrastructural in- 
vestigations using transmission electron 
microscopy (Quicke et al. 1992, Newman 
and Quicke 1999a), but the data obtained 
are not normally easily interpreted in 
terms of the actual size and structure of 
the spermatodesm. However, it was ap- 
parent that spermatodesmata vary in both 
size (i.e. number of individual sperm in- 
volved) and shape. For example, in most 
taxa examined the spermatodesmata re- 
semble a tuft of grass with their acrosomes 
embedded in an extracellular cap and the 
nuclei and tails splaying out posteriorly. 
However in the cephid, Ccphus pi/gmaeus, 
the whole spermatodesm is xery elongate, 
several times longer than an individual 
sperm, and sperm are iiiserted along a 
thin central extracellular matrix core 
(Quicke et al. 1992). 

Sperm produced in a given follicle are 
all derived from cell divisions from a sin- 



120 



Journal of Hymenoptera Research 



gle spermatogonia! cell (see Quicke 1997). 
Because these cell divisions occur synchro- 
nously within a follicle and each sperm 
mother cell in each follicle undergoes a 
fixed number of cell division rounds, the 
numbers of sperm per spermatodesm are 
expected to be 2" where n is the number 
of rounds of spermatocyte division. 

Counting the numbers of sperm in each 
spermatodesm was not straight-forward 
for most taxa because when stained using 
traditional dyes, the mass was so opaque 
with overlapping nuclei and tails that in- 
dividual cells could not be distinguished 
and counted. We have therefore employed 
a fluorescence staining technique in order 
to measure the total DNA content of the 
spermatodesm and divided that by the 
DNA content of an individual sperm nu- 
cleus. For a few taxa that were no longer 
available for the current study, we have 
included some crude estimates of sperm 
number obtained from transmission elec- 
tron microscopy of transverse sections 
(Quicke et al. 1992, Newman and Quicke 
1999a). However, these are likely to be un- 
derestimated, because more posteriorly in- 
serted sperm may not have been sec- 
tioned. 

MATERIALS AND METHODS 

Materials. — The following taxa were ex- 
amined. Xyelidae: Xifcla sp., Colorado; 
Tenthredinidae: Tciithrcdo xaiitlia Norton, 
N California; Stwngi/lo^asfcr disfaiis Nor- 
ton, S California; Dolcnis tcjonicnsis (Nor- 
ton), S California; Pergidae: Aconiulccera 
sp., Illinois; Cimbicidae: Trichiosoiiia triaii- 
gulujji (Kirby), NW California; Cimhcx 
amcricamim Leach, NW California; Anax- 
yelidae: Syniexis Uboccdrii Rohwer, Califor- 
nia; Xiphydriidae: Xipln/dria abdoniiualis 
Say, Illinois; Xiphydria maculata Say, Illi- 
nois; Orussidae: Onissus thoracicus (Ash- 
mead), N California; Orussus occideiitalis 
(Cresson), N California. It should be noted 
that males of many sawflies are taxonom- 
ically difficult to segregate and at present 
species-level identifications are not always 



possible. Voucher specimens have there- 
fore been deposited in the United States 
National Museum (Washington D.C.). 

Light microscopy. — Vas deferentia and 
testes were dissected from living sawflies 
in insect saline (Clark et al. 1979) and 
teased apart on a clean microscope slide. 
After a few minutes to allow the sperm/ 
spermatodesmata to swim free of the dis- 
rupted tissue, the slides were heat fixed on 
an hot plate at approximately 80°C. These 
slides were stained with a 0.007mg/ml so- 
lution of 4',6-diamidino-2-phenylindole 
dihydrochloride (DAPI) and viewed with 
a Leitz epifluorescence microscope at X 
1000 (as in Flemming et al. 2000). This 
stain specifically binds to double stranded 
DNA and fluoresces under UV light (Figs. 
1, 2). Images of stained spermatodesmata 
were captured using a CV-M300 video 
camera, and a Scion LG3 frame-grabber 
mounted in a Power Macintosh running 
Scion Image 1.62a. Care was taken to en- 
sure that, in each frame, both a complete 
spermatodesm and an isolated sperni cell 
was present (This was not possible for 
Acorduleccra where only c^ne isolated 
sperm nucleus could be found). Before 
capturing, the iniage was adjusted to min- 
imise pixel saturation and these adjust- 
ments affected the spermatodesm and iso- 
lated sperm cell in each image equivalent- 
ly. A densitometric value for both the 
spermatodesm and the sperm nucleus was 
determined (the product of area measured 
in pixels and staining intensity) and the 
nun"iber of sperm in the spermatodesm de- 
rived by dividing the two values. On av- 
erage four images per species were ana- 
lysed. For one species, Xiphydria iiiaciihita, 
it was possible to make a direct count of 
sperm nuclei present in the spermato- 
desm. Comparison of this value with that 
obtained densitometrically (35 ± 7 {95% 
confidence interval) and 33 ± 3 {95% con- 
fidence interval) respectively) confirms the 
accuracy of the technique. Basic statistics 
were calculated using Excel 98 (Microsoft). 

Making the assumption that DNA 



Volume 10, Number 2, 2001 



121 




Fig. 1. Fluorescence images of DAPI-stained sperm nuclei in spermatodesmata of sawflies investigated: A, 
Acordiilcccra sp. (Pergidae); B, Trichiosoiiia triaugului}i (Cimbicidae); C, Stnmgi/logastcr distaiis (Tenthredinidae); 
D, Dolcrus tcjouicusis (Tenthredinidae); E, Tciitlurdo xniitlia (Tenthredinidae); F, Cindvx americn)ium (Cimbici- 
dae). 



(chromatin) is densely packed in sperm 
nuclei, and since we have no n priori rea- 
sons to expect differences in DNA density 
between taxa, we used sperm nucleus size 
as a surrogate for haploid DNA content. 

RESULTS 

The sperm heads are inserted through- 
out the cap of the spermatodesmata, with 
those sperm located more centrally being 
inserted more anteriorly (Figs. 1, 2). For 
Xyelidae, Anaxyelidae, Xiphydriidae and 
Orussidae, the spermatodesmata are elon- 
gate structures, in the case of Orussus, the 
sperm appear to be inserted in the cap in 
a spiral configuration rather as if a cylin- 
drical roll of paper was 'pulled out' from 



the middle. Tenthredinoid spermatodes- 
mata are far wider and the larger ones (i.e. 
those with a larger number of sperm; Fig. 
la,b,f) dry on to the slides as rosette like 
structures. Although the centres of these 
appear empty in DAPI-stained material, 
transmission electron micrographs (New- 
man and Quicke 1999a) suggest that this 
is the region where the acrosomes are in- 
serted in the extracellular matrix of the 
cap. 

Results of numbers of sperm per spcr- 
matodesm are shown graphically in Fig. 3. 
Visual inspection of numbers (which as 
explained above are expected to be integer 
powers of 2) suggests that for Xi/cla, the 
number is 16, for Orussidae, Anaxyelidae 



122 



Journal of Hymenoptera Research 




Fig. 2. Fluorescence images of DAPI-stained sperm nuclei in spermatodesmata of sawflies investigated: A, 
Xi/t'/rt (Xyelidae); B, Si/ntcxis liboccdrii (Anaxyelidae); C, Xipln/drin abdonii)mlis (Xiphydriidae); D, Xiphydrin uia- 
ciilata (Xiphydriidae); E, Orussiis occidcntalis (Orussidae); F, Orussiis tlioracicuf (Orussidae). 



700 
i 600 
I 500 
^ 400 
t 300 
I 200 

u 

D 

^100 




sperm per Spermatodesm 

(± 95% Confidence Interval) 



.^^^ 





■ 


" 


p 


- 


T ^ ^ 


L 


ti 


'- -^ 


'- 






b 




, -'.'^'.'^"■.'^.'''^.riii 





^-^^ 



.o>'' 



Fig. 3. Plot of numbers of sperm per spermatodesm for sawfly taxa ranked according to spermatodesm size. 



Volume 10, Number 2, 2001 



123 



CU 1 



Mean Nuclear Size 

(± 95%Confidence Intej-val) 



1400 



1200 



000 
800 
600 
400 
200 -H 







.-rii— -^ 



A 



><■■ 



->^ 



^^ 



Fig. 4. Plot of mean nucleus size, a surrogate for DNA content, for indi\ idual sperm in each sawtly taxon. 



and Xiphydriidae, it is 32, for Tenthredi- 
nidae the number ranges from 128 to 256, 
for Pergidae 64 and for Cimbicidae, 256- 
512. Visual inspection of toluidine blue 
stained slides of two other tenthredinids 
{Rhogognstcr cnliforuicn (Norton) and a Teii- 
thredo that was either T. lacticiiicta Cresson 
or T. varipicta Norton) suggests that they 
have the same number of sperm per sper- 
matodesm as T. xnuiha. 

Counts of the number of pixels occu- 
pied by isolated sperm nuclei (Fig. 4) al- 
low us to estimate nuclear DNA content. 
The distribution is suggestive of a trend 
among the sawflies in that Xydn, the most 
basal genus, has the largest nucleus and 
Oriissiis, the most derived genus has the 
smallest nucleus. 

Although we have not been able to uti- 
lise the present technique to quantify sper- 
matodesm size in the Cephoidea or Siri- 
cidae, inspection of the stained light mi- 
crographs and of transmission electron 
micrographs for these two superfamilies 
respectively (Quicke et al. 1992, Newman 
and Quicke 1999a) indicate that both of 



these have rather large numbers of sperm 
per sperniatodesm. Our best estimates 
were taken as the smallest power of two 
larger than the definite minimum number 
of individual sperm within a micrograph 
of a spermatodesm. The sperm tails are 
relatively straight for a distance after 
emerging from the sperm head and in our 
micrographs the spermatodesmata were 
reasonably isolated so we do not believe 
that there is any reason why that sperm 
number will have been over-estimated. 
For the cephid, Cephiis, this is 128 (based 
on a count of 93 sperm tails), and for the 
siricid, Tremex, it is 512 (based on a count 
of c. 400 transverse sections of sperm in a 
largely complete micrograph section). 

DISCUSSION 

Given that we now ha\'e a robust phy- 
logenetic hypothesis for the superfamilies 
of sawflies and at least an estimate of fam- 
ily level relationships within the tenthre- 
dinoid lineage (Vilhelmsen 1997, 2000a,b, 
2001), we can consider the evolutionary 
pattern of spermatodesm size (number of 



124 



Journal of Hymenoptera Research 



5a 



15 Xyela sp. 
128 Tenthredo xantha 
256 Strongylogaster distans 
128 Dolerus tejoniensis 
256 Cimbex americanum 
512 Trichiosoma triangulum 
64 Acordulecera sp. 
32 Syntexis libocedrii 
32 Xiphydria abdominalis 
32 Xiphydria maculata 
32 Orussus occidentalis 
32 Orussus thoracicus 



5b 



15 Xyela sp. 

128 Tenthredo xantha 

256 Strongylogaster distans 

128 Dolerus tejoniensis 

256 Cimbex americanum 

512 Trichiosoma triangulum 

64 Acordulecera sp. 

.^72^ Cephus 

32 Syntexis libecedrii 

7512 Tremex 

32 Xiphydria abdominalis 

32 Xiphydria maculata, 

32 Orussus occidentalis 

32 Orussus thoracicus 



Fig. 5. Number of sperm per spermatodesm shown on tlie independently obtained cladogram of sawfly 
relationships (from Vilhelmsen 1997, 2001), showing in (a) only data obtained from DAPI-staining, and (b) 
with values for additional taxa based on other estimation techniques incorporated. 



sperm included). Unfortunntely, there is 
considerable uncertainty about what con- 
stitutes a suitable outgroup for the Hy- 
menoptera, and if one accepts a currently 
common view that the order is the sister 
group of the remainder of the Holometa- 
bola, then there is too much variation with 
this putative sister group to use it as an 
outgroup for the purposes of the current 



analyses. Therefore we base our interpre- 
tations on the likely ancestral state in the 
order on the state shown bv the Xyelidae 
which display the most putatively ple- 
siomorphic character states of any of the 
extant Hymenoptera. Visual inspection of 
the DAPI sperm-equantification data 
mapped on to Vilhelmsen's {loc. cit.) in- 
dependently derived sawfly phylogeny 



Volume 10, Number 2, 2001 



125 



(Fig. 5a) suggests that the groundplan 
spermatodesm size for sawl^ies is low (16 
or 32), but that there has been a general 
increase within the Tenthredinoidea 
(range 64-512) and that particularly large 
numbers (256-512) hav^e evolved at least 
twice within this superfamily. However, 
incorporating estimates of sperm number 
for Ccphus pyguiciis (Cephoidea) and Trc- 
mcx sp. (Siricidae) into Vilhelmsen's phy- 
logeny [loc. cit.) (Fig. 5b) tends to confuse 
the picture in that it is equally parsinio- 
nious that there was a marked increase in 
sperm number per spermatodesm above 
the Xyelidae, and that there were reversals 
to lower numbers in the Anaxyelidae and 
Xiphydriidae + Orussidae as for multiple 
increases from a groundplan of 16 or 32 
(viz in the Tenthredinoidea, Cephoidea 
and Siricidae). 

Denser taxon sampling may help to 
clarify the above issues and may also pro- 
vide additional phylogenetic evidence 
within some groups, especially within the 
Tenthredinoidea. Future work will exani- 
ine nuclear DNA content across the Hy- 
menoptera in more detail (Schiff, Flem- 
ming and Quicke in preparation). 

ACKNOWLEDGEMENTS 

We would like to thank Virginia Scott for sending 
us the li\'e male Xycla, David Smith for comparing 
the Teiithrcdo voucher specimens with the USNM 
(Washington DC) reference collection, and Larry Be- 
zark and Steve Heydon for helping and providing NS 
with company on some of the collecting trips. 

LITERATURE CITED 

Clark, R. B., K. A. F. Gration and P. N. R. Usherwood. 
1979. Desensitization of glutamate receptors cmt 
innervated and denervated locust muscle fibres. 
Journal of Plnjsiologi/ (London) 290: 551-568. 

Flemming, A. J., Z. Z. Shen, A. Cunha, S. U. Emmons 
and A. M. Leroi. 2000. Somatic polyploidisation 
and cellular proliferation drive body size evolu- 
tion in nematodes. Proceedings of the Niitlonnl 
Academy of Sciences, USA, 97: 5285-5290. 

Jamieson, B. G. M. 1987. Tlie Ultrastructure and Pln/- 
logon/ of hisect Spermatozoa. Cambridge Unix'er- 
sity Press, Cambridge, 320pp. 



Lino-Neto, J., S. N. Bao, and H. Dolder. 1999. Struc- 
ture and ultrastructure of the spermatozoa of Be- 
plvatelloides poniorum (Fabricius) (Fiymenoptera: 
Eurytomidae). International journal of bisect Mor- 
phologif and Embri/ologi/ 28: (4) 253-259. 

Lino-Neto, J., S. N. Bao, and H. Dolder. 2000a. Struc- 
ture and ultrastructure of the spermatozoa of Tri- 
chogrnmma pretiosum Riley and Trichograuuua ato- 
povirilia Oatman and Platner (Hymenoptera: Tri- 
chogrammatidae). Acta Zooloogica, Stockholm 81: 
205-211. 

Lino-Neto, J., S. N. Bao, and H. Dolder. 2000b. Sperm 
ultrastructure of the honey bee {Apis mellifera) (L) 
(Hymenoptera, Apidae) with emphasis on the 
nucleus-flagellum transition region. Tissue and 
Cell 32: 322-327. 

Newman, T. M. and D. L. ]. Quicke. 1998. Sperm de- 
velopment in the imaginal testes of Aleiodes cox- 
nlis (Hymenoptera: Braconidae: Rogadinae). /oin- 
nal of Hi/Dienoptera Research 7: 25-37. 

Newman, T. M. and D. L. J. Quicke. 1999a. Ultrastruc- 
ture of imaginal spermatozoa of sawflies (Insec- 
ta: Hymenoptera: Symphvta). Journal of Hi/me- 
noptera Research 8: 35-47. 

Newman, T. M. and D. L. J. Quicke. 1999b. Ultra- 
structure of spermatozoa in Leptopilina (Hvme- 
noptera: Cynipoidea: Eucoilidae). journal of Hy- 
}uenoptera Research 8: 197-203. 

Newman, T. M. and D. L. J. Quicke. 2000. Sperm de- 
velopment and ultrastructure of mature sper- 
matozoa of Megahfra (Hymenoptera: Megalyro- 
idea). ]ournal of Hi/menoptera Research 9: 62-70. 

Quicke, D. L, J, 1997. Parasitic Wasps. Chapman & 
Hall, London, 470pp. 

Quicke, D. L. J., S. N. Ingram, H. S. Baillie and P. V. 
Gaitens. 1992. Sperm structure and ultrastructure 
in the Hymenoptera (Insecta). Zoologica Scripta 
21: 381-402. 

Vilhelmsen, L. 1997. The phylogeny of lower Hyme- 
noptera (Insecta), with a summary of the early 
evolutionary history of the order. Journal of Zoo- 
logical Si/stematics ami Evolutionary Research 35: 
49-70. 

Vilhelmsen L. 2000a. Before the wasp-waist: Com- 
parative anatomy and phylogenetic implications 
of the skeleto-musculature of the thoraco-abdom- 
inal boundary region in basal Hymenoptera (In- 
secta). Zoomorphologif 119: 185-221. 

Vilhelmsen L. 2000b. The ovipositor apparatus of bas- 
al Hymenoptera (Insecta): phylogenetic implica- 
tions and functional morpholog}'. Zoologica Scrip- 
ta 29; 319-345. 

Vilhelmsen L. 2001. Phylogeny and classification of 
the extant basal lineages of the Hymenoptera (In- 
secta). Zoological journal of the Linnean Society 131: 
393^142. 



J. HYM. RES. 
Vol. 10(2), 2001, pp. 126-130 

First Biological Data for Aspilodemon Fischer (Hymenoptera: 
Braconidae: Hydrangeocolinae): Parasitoids of Cecidomyiid Fly Galls 

on Asteraceae in Brazil 

Rubens A. M. Oda, Margarete Valverde de Macedo, and Donald L. J. Quicke 

(RAMO, MVM) Lahoratorio de Ecologia de Insetos, Departamento de Ecologia, CP 68020, 
Instituto de Biologia, Universidade Federal do Rio de Janeiro, Rio de Janeiro CEP 21941-590, 

Brazil; (DLJQ) Unit of Parasitoid Systematics, Department of Biology, Imperial College at 

Silwood Park, Ascot, Berkshire SL5 7PY, U.K., and Department of Entomology, The Natural 

History Museum, London SW7 5BD, UK 



Abstract. — Biological data for the hydrangeocoline braconid genus Aspilodemon Fischer are re- 
porteti for the first time. An Aspilodefuoii sp. from Brazil is reported as a parasitoid of three species 
of Liodiplosis spp. (Diptera: Cecidomyiidae) which induce galls on leaves of the liana, Mikaiiin 
glouwrata (Asteraceae); it does not attack other gall-forming cecidomyiids on the same plant, and 
evidence suggests that it is ectoparasitic although direct observations of parasitoid larvae have 
not been made. How this affects our understanding of the evolution of gall formatic^n and aphid 
parasitism by braconids is discussed. 



The l-iydrangeocolinae are a recently 
recognised subfamily of braconid wasps 
(Dowton et al. in press, Belshaw and 
Quicke in press), to date known from 
three genera, each with a southern ciistri- 
bution, viz Hi/dniiigcocohi Brethes ( = Ke- 
pJmloscDia Fischer) from Chile, Aspilodeiuoii 
Fischer, from Central and South America 
and Opioptcnis Szepligeti (see Wharton 
1993, Whitfield and Wharton 1997), from 
Australia. Until recently, some of these 
taxa were typically treated under the sub- 
family Opiinae (e.g. Fischer 1966, but see 
Wharton 1988) or more recently the Hor- 
miinae, see e.g. Whitfield and Wharton 
(1997) or Rhyssalinae (Quicke et al. 1997) 
though several of these authors recog- 
nised, they do not fit well into any of these 
(see electronic appendix of Belshaw et al. 
2000 for further discussit^n). Interest in the 
group was recently increased because mo- 
lecular sequence data strongly indicate 
that they are not related to Opiinae, Ht^r- 
miinae or Rhyssalinae, but instead form a 
sister group to the endemic Australian 
subfamily, Mesostoinae, and that together 



these are the sister group of the well 
known Aphidiinae (Belshaw et al. 2000). 
This newly discovered relationship was 
particularly interesting as it appeared to 
suggest both a Gondwanan origin of the 
Aphidiinae despite their currently pre- 
dominantly northern distribution, and a 
link with galls, because the most primitive 
extant aphids are gall formers, mesostoine 
braconids form galls on Baiiksia spp. (Pro- 
teaceae) (Austin and Dangerfield 1998) 
and the only hydrangeocoline for which 
some biology is known, Hi/drniigcocola, has 
been reared from unidentified galls on Hi/- 
drniigca (Hydrangeaceae) (Brethes 1927). 
Very little is known, however, about the 
biologies of the putatively most primitive 
extant aphidiines, and a direct association 
of any these with gall-forming aphids is 
yet to be confirmed. 

Here we present the first rearing data 
for the genus Aspilodemon based on a 
probably undescribed species from Brazil. 
The species concerned is associated with 
cecidomyiid fly galls on Mikniiia glomcnitn 
Sprengel, and the available evidence 



Volume 10, Number 2, 2001 



127 



Table 1. Number of galls collected in each locality during study; gall types are illustrated in Gagne et al. 
(2001). 









Morpholoj 


jical type of gall 








Locality 


Cylindrical 


Spherical 


Conical 


Leaf/Vein 


Bi.cl 


Epidermis 


Stem 


Parati 


2070 


1085 





885 


5 


53 


27 


Itatiaia 


1385 


468 





574 


11 


17 


22 


Po^x-. 


1061 


583 


17 


272 


5 


17 





Picinguaba 


861 


471 


188 


305 


2 


38 


9 


S. Orgaos 


297 


361 





252 


2 


20 


56 


Tijuca 





6 


824 


51 





141 






strongly suggests that it is a parasitoid of 
the tly larva. M. glouiernta is a liana species 
belonging to the Asteraceae, and it occurs 
inside and on the edges of forests, flow- 
ering from August to December. The ge- 
nus Mlkniiia has 415 species mainly dis- 
tributed in Central and South America, of 
which 171 species occur in Brazil (King 
and Robinson 1987). M. gloiucrata ranges 
from north-eastern Brazil down to the 
southern-most part of Brazil and just into 
Argentina and Paraguay (Ritter et al. 
1992). 

MATERIALS 

Voucher specimens of the Aspilodcmou 
sp. reared are deposited in each of the fol- 
lowing collections: The Natural History 
Museum, London; the Entomological Col- 
lection of the Laboratorio de Ecologia de 
Insetos, Department of Ecology, Univer- 
sidade Federal do Rio de Janeiro. Morpho- 
logical terminology follows Sharkey and 
Wharton (1997). The D2-D3 28S rDNA 
gene sequence for the species of Aspllodc- 
mon referred to here is deposited in EMBL: 
accession number AJ245685 (and has been 
used in phylogenetic analyses of Belshaw 
et al. 2000 and Quicke and Belshaw 1999). 

Study areas and sampling. — The research 
was based in the Atlantic coast (Mata At- 
lantica) forests of Brazil which has one of 
the highest levels of biological diversity in 
the world, and is representative of humid 
tropical forests and their associated eco- 
systems (Mori et al. 1981). The localities 
were (1) Itatiaia National Park, Itatiaia 



County, Rio de Janeiro State; (2) Serra dos 
Orgaos National Park, Teresopolis Coun- 
ty, Rio de Janeiro State; (3) Tijuca National 
Park, Rio de Janeiro County, Rio de Janei- 
ro State; (4) Parati County, Rio de Janeiro 
State; (5) Picinguaba State Park, Ubatuba 
County, Sao Paulo State; (6) Biological Re- 
serve of Po(;o das Antas, Silvia Jardim 
County, Rio de Janeiro State. Four collec- 
tions of galls (one per season — one day in 
the field) were made at each of the first 
five of the above localities: April or May, 
1998 (Autumn), July or August 1998 (Win- 
ter), October or November 1998 (Spring) 
and January or February 1999 (Stimmer). 
The sixth locality was sampled at monthly 
intervals from February 1996 to October 
1997. Total numbers of each type of gall 
collected at each field site are given in Ta- 
ble 1. 

Eight species of cecidomyiid flies form 
galls on Mikania glonierata in Brazil, and 
the galls of most of these can be distin- 
guished on the basis of their morphology 
and location on the plant (Table 2) (see 
Gagne et al. 2001, in which gall types are 
also illustrated). Cecidomyiid galls were 
collected from M. glomcrata plants at six 
localities and reared to discover what par- 
asitoids might be attacking them. Some 
galls were dissected. 

OBSERVATIONS 

Rearings of Aspilodemon. — Rearings are 
summarized in Table 3. The numbers of 
each type of gall collected at each locality 
(Table 1) are estimates of their relative 



128 



Journal of Hymenoptera Research 



Table 2. Cecidoinyiid fly galls found on Mikania 
glomemia (Asteraceae) with descriptions of gall loca- 
tion and type. 

Super- tribe Cecidomyiidi 

Mikniiiadip>losis niuiulipcs (Gagne) — leaf vein 
swelling 
Tribe Clinodiplosini 
Liodiplosis ci/li)h1rica Gagne — cylindrical leaf 

gall 
L. conica Gagne — conical leaf gall 
L. sphcrica Gagne — spherical leaf gall 
Tribe Asphondyliini 
AsphoiuiyUn gloincmtnc Gagne — \'ein swelling 
A. luochiiii Skurava — stem swelling 
Pcmsplioiidi/lm iiiikaiiine Gagne — bud gall 
Super-tribe Lasiopteridi 
Tribe Alycaulini 
Ah/caiihis globulus Gagne — leaf epidermis 
swelling 



abundance, and therefore it is clear that 
although the cylindrical gall type was 
niore abundant by approximately a factor 
of 2 at the three localities from where most 
individuals of Asplloiicinoii were obtained 
(Parati, Picinguaba and Pocjo), the Aspnlo- 
dojioii demonstrated a clear preference for 
the spherical gall type which yielded pro- 
portionately far more wasps (Table 3). 

Except for the only individual obtained 
from the leaf vein swelling or vein swell- 
ing (we could not differentiate these two 
gall types) at Picinguaba, all Aspiloilcnioii 
individual were obtained from the galls 
induced by species of Liodiplosis. As we 
collected and reared many galls it is pos- 
sible that the individual supposed to have 
been obtained from leaf vein swelling or 
vein swelling is a contamination. 

Numerous Chalcidoidea were reared 
from the three gall types attacked by As- 
pilodeinon; at least 10 species from the con- 
ical leaf gall type alone (taking all locali- 
ties and sampling dates into account). The 
importance of Aspilodcmon as a parasitoid 
(in terms of parasitism rate) varied: it was 
the commonest parasitoid reared from the 
spherical and conical leaf galls and the 
fourth commonest in the cylindrical gall. 

Evidence that Aspilodemon is a parasitoid 



Table 3. Total Aspilodemon rearings from cecido- 
myiid galls on Mikania gkvnerata at each site. 



Morphological type of g.il 



Localit\ 



C\lindiK\il Sphenc.il Conical Lcat/Vein 



Parati 




19 


65 





Itatiaia 




1 








Voc^o 




2 


44 


3 


Picingu 


aba 


4 


9 


2 


S. Orgaos 





2 






and not a gall-fornicr. — Aspilodemon individ- 
uals were always found inside Cecido- 
myiidae gall chambers, and all the types 
of gall found (Table 1) yielded cecido- 
myiid flies showing that no gall type 
yielded only Aspilodemon and no potential 
hosts. Further, the Aspilodemon species 
was associated with three morphological- 
ly different gall types which would not be 
expected if it was a gall inducer itself. 

Evidence that Aspilodemon mai/ be ecto- 
parasitic. — Aspilodemon pupates inside a 
cocoon which occupies approximately half 
of the cecidomyid chamber. All the endo- 
parasitoids that were also reared (see 
above) pupated inside the cecidomyid 
skin and we could find the cecidomyid 
larval sternal spatula (a strongly scleroti- 
sed thoracic feature) within the galls after 
these endoparasitoids had emerged. When 
Aspilodemon parasitised the cecidomyiid 
no host remains could be found, perhaps 
indicating that the sternal spatula had 
been consumed. 

DISCUSSION 

The discovery that Aspilodemon is asso- 
ciated with galls strengthens the possibil- 
ity that this way of life is the norm for the 
subfamily Hydrangeocc^linae. Previously 
published suggestions that Aspilodemon 
belongs to the Opiinae (Fischer 1966) have 
been superseded by both morphological 
(Wharton 1988, Whitfield 1993) and mo- 
lecular phylogenetic analyses (Belshaw et 
al. 2000), which instead support a relation- 
ship with the Australian subfamily Me- 
sostoinae and also with the Aphidiinae. 



Volume 10, Number 2, 2001 



129 



Several transitions to cecidogenesis 
have occurred in the Braconidae: Mesos- 
toinae appear to be exclusively cecidogen- 
ic on Bimksla species (Proteaceae) (Quicke 
and Huddleston 1989, Austin and Dan- 
gerfield 1998); the enigmatic genus Moni- 
toriclhi Hedqvist produces galls on Philo- 
dendroii (Liliaceae) (Infante et al. 1995); 
several species of the doryctine genus Al- 
lorlio^^ns Gahan (Macedo and Monteiro, 
1989; Marsh et al. 2000), and another do- 
ryctine, Pscnobolus, is an inquiline in figs 
(Ramirez and Marsh 1996) and may be 
partly cecidogenic. Knowledge that at 
least some Hydrangeocolinae (precise bi- 
ology of Hi/dnmgeocohi is still unknown) 
are specialist parasitoids of gall-forming 
Diptera while the closely related Mesos- 
toinae are cecidogenic suggests that the 
latter biology could have evolved from the 
former, as has been suggested for cecido- 
genic Eurytomidae (Chalcidoidea). It 
would be interesting to know whether any 
hydrangeocolines have also made this 
transition. 

We wish to emphasise that we have not 
yet obtained any strong evidence about 
whether Aspilodeuioii is ecto- or endopar- 
asitic. Although no host remains were 
found in galls parasitised by this wasp, in- 
dicating that all the host cuticle had been 
consumed, it should be noted that most 
endoparasitic braconids (those belonging 
to the 'helconoid' and 'n"iicrogastroid' 
groups of subfamilies — see Belshaw et al. 
2000), very often emerge from their hosts 
before pupation and have an external 
feeding phase (Shaw and Huddleston 
1991, Shaw and Quicke 2000). So even if 
the sternal spatula has been consumed it 
does not necessarily mean that the para- 
sitoid was not endoparasitic in its earlier 
instars. 

Aspilodciiioii differs frc^m Hi/dnmgcocoln 
in only one fixed character — the fore wing 
pterostigma is narrow but distinct from 
vein Rl in the latter, whereas there is nc^ 
discernible pterostigma in the former. In 
addition, Aspilodeuiou always lacks fore 



wing vein 2a, whereas it is present in most 
Hi/draugeocola. Given that for both these 
characters, Hydraugeocoln displays (at least 
most species) the putatively plesiomorph- 
ic state, it is likely that the species classi- 
fied under Aspilodeuiou may simply be a 
derived group of species within a broader 
concept of Hi/drnugcocola. 

ACKNOWLEDGEMENTS 

We would like to acknowledge Ricardo F. Montei- 
ro (UFRJ) for helping during all phases of the work, 
Raymond Gagne for identifying and describing the 
Cecidomyiidae species, IBAMA and lEF/SP for au- 
thorization to collect in the Units of Conservation, 
CNPq for the scholarships to R.A.M. Oda and finan- 
cial support. 

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Belshaw, R., M. Dowton, D. L. J. Quicke, and A. D. 
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Belshaw, R., E. Herniou, C. Gimeno, M. G. Fitton and 
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Brethes, ]. 1927. Nouveaux Hymenopteres parasites 
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Dowton, M., R. Belshaw, A. D. .Austin, and D. L. J. 
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Fischer, M.1966. Aspilodenton, ein neues Opiinen-Ge- 
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Journal of Hymenoptera Research 



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Infante, F., P. Hanson and R. A. Wharton. 1995. Phy- 
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Macedo, M. V. de and R. F. Monteiro. 1989. Seed pre- 
dation by a braconid wasp, Allorliogas sp. (Hy- 
menoptera). Journal of the New York Entomological 
Society 97: 358-362. 

Marsh, P. M., M. V. de Macedo and .M. C. P. Pimentai. 
2000. Descriptions and biological notes on two 
new phytophagous species of the genus Allorlio- 
gas from Brazil (Hymenoptera: Braconidae: Do- 
ryctinae). journal of Hymenoptera Research 9: 292- 
297. 

Mori, S. A., B. M. Boom and G. T. Prance. 1981. Dis- 
tribution patterns and conservation of eastern 
Brazilian coastal forest species. Brittonia 33: 233- 
245. 

Quicke, D. L. J., C. van Achterberg and H. C. J. God- 
fray. 1997. Comparative morphology of the ven- 
om gland and reservoir in opiine and alysiine 
braconid wasps (Insecta, Hymenoptera, Bracon- 
idae). Zoologica Scripta 26: 23-50. 

Quicke, D. L. J. and R. Belshaw. 1999. Incongruence 
between morphological data sets: an example 
from the evolution of endoparasitism among 
parasitic wasps (Hymenoptera: Braconidae). Sys- 
tematic Biology 48: 436-454. 

Quicke, D. L. J. and T. Huddleston. 1989. The Aus- 



tralian braconid wasp subfamily Mesostoinae 
(Hymenoptera: Braconidae) with the description 
of a new species of Mesostoa. journal of Natural 
Historx/ 23: 1309-1317. 

Ramirez, W. B. and P. M. Marsh. 1996. A review of 
the genus Psenobolus (Hymenoptera: Braconidae) 
from Costa Rica, an inquiline fig wasp with bra- 
chypterous males, with description of two new 
species. Journal of Hymenoptera Research 5:64-72. 

Ritter, M. R., L. R. M. Baptista, and N. I. Matzen- 
bacher. 1992. Asteraceae genero Mikania Willd 
Sec(;6es Globosae e Thirsigerae. Boletim do Insti- 
tuto de Biociencias / UFRGS 50: 1-90. 

Sharkey, M. J. and R. A. Wharton. 1997. Morphology 
and terminology. In R. A. Wharton, P. M. Marsh 
& M. J. Sharkey (eds) Identification manual to the 
Neiv World genera of Braconidae. Special Publica- 
tion of the International Society of Hymenopter- 
ists 1: 19-37. 

Shaw, M. R. and T. Huddleston. 1991. Classification 
and biology of braconid wasps (Hymenoptera: 
Braconidae). Handbooks for the Identification of Brit- 
ish Insects 7(U): 1-126.. 

Shaw, M. R. and D. L. J. Quicke. 2000. The biology 
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Natural History 34: 611-628. 

Wharton, R. A. 1988. Classification of the braconid 
subfamily Opiinae (Hymenoptera). The Canadian 
Entomologist 120:333-360. 

Wharton, R. A. 1993. Review of the Hormiini (Hy- 
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Whitfield, J. B. and R. A. Wharton. 1997. Subfamily 
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285-301. 



J. HYM. RES. 
Vol. 10(2), 2001, pp. 131-137 

First Record of Aridehis nifotestaceus Tobias (Hymenoptera: 

Braconidae, Euphorinae) Parasitizing Nezara viridula Nymphs 

(Heteroptera: Pentatomidae) with Observations on its Immature 

Stages and Development 

Scott R. Shaw, Gianandrea Salerno, Stefano Colazza, and Ezio Peri 

(SRS) Insect Museum, Department of Renewable Resources, University of Wyoming, Laramie, 

WY 82071-3354, USA; (GS) Department of Arboriculture and Plant Protection, University of 

Perugia, 06122 Perugia, Italy; (SC, EP) Institute of Agricultural Entomology, University of 

Palermo, Palermo, Italy 



Abstract. — Aridehis nifotestaceus Tobias is recorded for the first time from Italy as parasitizing 
the pentatomid bug Nezara viridula L. This is the first record of the species in southern Europe 
and the first host data. The species is re-described and illustrated. New information is provided 
on its immature stages, development, and biological control potential. 



Species of the euphorine braconid genus 
Aridehis Marshall are cosmopolitan in dis- 
tributit^n, but most diversified in tropical 
areas (Shaw 1985). Aridehis species are 
cjuite distinctive in appearance, and can be 
easily distinguished from other braconids 
by their coarse honey-combed areolate 
mesosomal sculpture in combination with 
the long, tubular first metasomal segment 
and fore wing with a closed second sub- 
marginal cell (Shaw 1997). Their biology 
is not well known but the available re- 
cords indicate that they are solitary koino- 
biont endoparasitoids of heteropteran 
bugs in the families Pentatomidae, Platas- 
pidae, Scutelleridae, and Acanthosomati- 
dae (Kirkpatrick 1937; Shenefelt 1969; 
Papp 1974; Capek and Davidova-Vilimova 
1978; Tobias 1986; Shaw 1988; Maeto and 
Kudo 1992). 

Papp (1965) provided a taxonomic 
monograph of the world species of Aride- 
his, however, six Afrotropical species de- 
scribed by De Saeger (1946) were not in- 
cluded in Papp's monograph. Later, Papp 
(1974) erected the genus Arideloides for a 
species from New Guinea, but Shaw 
(1985) transferred the species to Aridehis. 



He (1980) describeci a new species from 
China, Chou (1987) revised the species of 
Taiwan, and, most recently, Chen and van 
Achterberg (1997) revised \he Arideliis spe- 
cies of China. They indicated that about 40 
Arideliis species are now known, of which 
20 are recorded from China. Shaw (1985) 
estimated that there are at least ten un- 
described Arideliis species in the Neotrop- 
ical region. Despite recent taxonomic 
work, until now only one species of Ari- 
deliis has been recorcied from Europe (She- 
nefelt 1969; Papp 1974; Capek and Davi- 
dova-Vilimova 1978). 

The purpose of this paper is to provide 
new host and distribution records for Ar- 
idehis riifotestnceiis Tobias recently discov- 
ered in Italy parasitizing the pentatomid 
bug Nezara viridula (L.). This is the first re- 
cord of the species in southern Europe and 
the first host data. The host, Nezara viri- 
dula, is one of the most serious agricultural 
insect pests worldwide, damaging a wide 
variety of fruit, nut, grain, and vegetable 
crops. It is the primary pest of soybean in 
many parts of the world (Todd 1989) and 
it also attacks many wild hosts that serve 
as reservoirs until agricultural crops are 



132 



Journal of Hymenoptera Research 



available (Jones and Sullivan 1988). In It- 
aly, annual losses due to this pest fluctu- 
ate in relation to changes in population 
abundance of N. viridiiln (Zandigiacomo 
1990; Colazza and Bin 1990, 1995). Al- 
though this pest has been the focus of nu- 
merous biological control programs, most 
of the recorded biological control agents 
attack the egg stage. The only other par- 
asitoids known to attack the nymphal and 
adult stages of N. viriduln are Tachinidae 
and Encyrtidae. Until now no species of 
Braconidae has been discovered attacking 
this host (Jones 1988). 

MATERIALS AND METHODS 

Periodically, during summer and au- 
tumn 1998 and 1999, adults and nymphs 
of N. viridula were collected in the fields 
in Umbria, Lazio and Sicily regions fol- 
lowing the seasonal sequence of host plant 
species. Most of the specimens were col- 
lected on maize and various vegetable 
plants. A study colony of the parasitoid 
was established and maintained in Italy 
by GS. Parasitoids were reared in the lab- 
oratory at a temperature of 24 plus or mi- 
nus 1 degree C, relative humidity 65'/o 
plus or minus 5%, and light/dark condi- 
tions of 16 hours light and 8 hours dark- 
ness daily. The insects were kept in plastic 
boxes and fed with vegetables and sun- 
flower seeds. Boxes were examined daily 
to collect parasitoid cocoons. Also, each 
day 5 N. viridula nymphs of the same age 
(reared in the laboratory) were exposed to 
a parasitoid female in a plastic box (7 X 
5.5 X 2.5 cm) for 24 h. The parasitized 
nymphs were removed from the box and 
kept separately until appearance of para- 
sitoid cocoons. 

Some nymphs were dissected to observe 
the different developmental stages of the 
parasitoid. For SEM analysis the immature 
stages were fixed in Karnovsky's medium 
(Karnovsky 1965) for one hour at 4 °C, de- 
hydrated in graded ethanol series, critical- 
point dried, mounted on stubs, coated 
with gold and observed with a Philips EM 



515 scanning electron microscope. Adult 
specimens were preserved in 95% ethanol 
and sent to SRS for description. Preserved 
adult specimens were transferred to 100% 
ethanol for 24 hours, then into chloroform 
for 30 minutes prior to drying and point- 
mounting to prevent shrinkage. 

Arideliis rufotestnceus Tobias, 1986 

■ (Figs. 1-10) 

Description of adult female. — length of 
body 4.8 mm; length of fore wing 3.5 mm. 
Head: Width of head in ciorsal view 2.1 
times its length; length of first tlagelk^m- 
ere 1.5 times length of second flagellom- 
ere; length of first and penultimate flagel- 
lomeres 4.5 and 0.8 times their width, re- 
spectively; median frontal carina weakly 
developed and somewhat obscured by 
coarse punctate sculpturing; vertex sculp- 
ture densely punctate; ocellar-ocular dis- 
tance 4.5 times ocellar diameter; occipital 
carina dorsally well-developed and com- 
plete; length of eye in dorsal view 1.6 
times length of temple; face and clypeus 
scultpture densely and coarsely punctate; 
intertentorial line 1.6 times length of ten- 
torial-ocular line; malar space 0.33 times 
height of eye. Mesosoiiia: 1.8 times longer 
than wide in dorsal view, densely areo- 
late. Wings: Pterostigma 2.0 times longer 
than wide at midpoint, anterior margin 
distinctly rounded and protruding well 
beyond anterior margin of wing as delim- 
ited by vein C-l-SC-(-R; length of marginal 
cell 0.85 times pterostigma length; vein r 
nearly perpendicular to pterostigma and 
2.5 times longer than vein 3RSa bordering 
second submarginal cell dorsally; vein 
3RSb nearly straight basally then curving 
towards wing margin apically; vein m-cu 
slightly antefurcal relative to vein 2RS, 
with very short segment of vein (RS + M)b 
present. Metasoiua: Entirely smooth and 
highly polished; length of first metasomal 
segment 7.0 times its width at spiracles; 
metasoma beyond petiole 2.7 times longer 
than wide in dorsal view; ovipositor 
sheath very short, exposed portion about 



Volume 10, Number 2, 2001 



133 




Figs. 1-5. Aridchi^ nttotc^Uucus. \, Eggs with developing embryos, 200 x. Figs. 2-4. First instar larva. 2, Lar\'a 
still partially surrounded by trophamnion and teratocytes, ventral view, 70x. 3, Lateral view, lOOx. 4, Ventral 
view of head capsule and mouthparts, 170x. 5, Third instar larva, antero-ventral view of head capsule and 
mouthparts, 105x. 



0.5 times length of hind basitarsus. Color. 
Head, antenna basally, lateral borders of 
pronotum, legs, and metasoma orangish 
brown; mandible apically, ocellar triangle, 



remainder of mesosoma, and ovipositor 
sheath black; wing venation brown, mem- 
brane clear to slightly dusky medially. 
Variation. — Body position at cieath vary- 



134 



Journal of Hymenoptera Research 




Figs. 6-10. Aridclus rnfotcstaceiis. 6, Third instar larva, ventral view, 11 X. Figs. 7-10 Adult. 7, Female near 
Neznrn viridiila nymph. 8, Propodeum and metasoma, dorsal view, 42x. 9, Propodeum, dorsal view, 38x. 10, 
Head, mesoscutum, and scutellum, dorsal \'iew, 42x. 



ing from metasoma fully extended poste- 
riorly to fully extended anteriorly (ovipo- 
sitional stance) with metasomal petiole 
bent under mesosoma and apex of meta- 
soma extending well beyond face. Sonie 
individuals appear darker with the head 



dorsally, hind femur, hind tibia, petiole, 
and dorsum of metasoma posteriorly 
more or less infused with sn^iokey black 
pigmentation. \n all cases dead preserved 
specimens appear somewhat to have a 
darker mesosoma; while alive some or- 



Volume 10, Number 2, 2001 



135 



angish brown color shows through the 
darker black pigmentation. Aside from 
genitalic differences, the male is quite sim- 
ilar in form and sculpture, but is much 
lighter in color appearing mostly orange, 
even over the mesosoma where black pig- 
mentation is limited to smokey pigmen- 
tation along the borders of the areolation. 
Description of iuunaturc stages. — The egg 
is alecithal (with no visible yolk), oval, 
with a clear chorion through which the 
white enibryo and developing tropham- 
nion are visible. The developing embryo 
has a large oval head capsule, followed by 
12 similar undifferentiated body seg- 
ments. The thoracic segments are not vis- 
ibly different from the abdominal seg- 
ments. The trophamnion forms a large 
mass of spongy white teratocytes below 
the embryo, enveloping the embryo pos- 
teriorly. The mature embryo has a thick 
round head capsule with no trace of eyes 
or antenna, deep anterior tentorial pits, 
long sickle-like mandibles, simple mouth 
opening, 11 undifferentiated similar body 
segments, and 12"' segment longer bearing 
anus ventrally and a long tapering caudal 
appendage. The first instar larva is of the 
caudate form, similar to the mature em- 
bryo with a thick round head capsule with 
no trace of eyes or antenna, deep anterior 
tentorial pits, long sickle-like mandibles, 
simple mouth opening, 11 undifferentiat- 
ed similar body segments, and 12* seg- 
nient k^nger bearing anus ventrally and a 
long tapering caudal appendage densely 
covered with short, thick, flexible setae. 
The first instar has an apneustic respira- 
tory system, with no visible spiracles. The 
body becomes much thicker as the young 
larva feeds and grows. The second instar 
larva becomes hymenopteriform remains 
apneustic. The sclerotized head capsule is 
much smaller, with short mandibles, and 
becomes enveloped by the fleshy first tho- 
racic segment as the larva grows. The sec- 
ond instar larva is yellowish white with 
undifferentiated segments, less distinct 
than in the first instar, and about 5x longer 



than wide. The caudal appendage is lost. 
The third and final iiistar larva is also hy- 
menopteriform and apneustic, but thicker 
and more maggot-like. It is tapering at 
both ends and thickest medially, being 
about 3x longer than wide at maturity. 

Biology. — Aridelus rufotestnceus was 
found for the first time in October 1998 
near the Umbria region of Perugia, Italy 
(parasitism rate 4.3%). During the summer 
of 1999 we found the parasitoid in the La- 
zio region (parasitism rate 21.7%) and in 
Sicily (parasitism rate 12.5%). The mature 
egg is usually lemon-shaped, with a ped- 
icel. When the first instar larva hatches 
from the egg, the teratocytes dissociate 
into the hemolymph and increase in size. 
In laboratory conditions the period from 
egg deposition to emergence of the mature 
larva was 23.18 plus or minus 2.77 days 
(n = 37). The mature larva emerges from 
the host through a hole in the interseg- 
mental membrane between the ultimate 
and penultimate segments, crawls away, 
and spins an oval white silk cocoon. After 
emergence of the parasitoid larva the host 
may survive for several days (although 
clearly not in healthy condition). Adults 
emerged from the cocoon in 22.27 plus or 
minus 1.45 days (n = 37). The adult life 
span was, in mean, 212.08 plus or minus 
8.18 days (n = 106) with a range from 6 
to 43 days. Reproduction is parthenoge- 
netic (thelyotokous, or sometimes deuter- 
otokous). In the lab only 3 males were ob- 
tained relative to 200 females. Prior to ovi- 
position females approach potential hosts 
on foot and inspect them, both visually 
and via antennation. Oviposition is typi- 
cally very rapid, lasting a few seconds at 
most, during which the female rapidly ap- 
proaches the host on foot, throws back the 
antennae, flexes the metasoma under the 
mesosoma while both exserting the ovi- 
positor and telescoping posterior metaso- 
mal segments. Eggs are inserted into the 
membranous cervical region between the 
head and thorax or into the intersegmental 
areas of the posterior abdominal region of 



136 



Journal of Hymenoptera Research 



the host. Supernumerary eggs of larvae 
were dissected from hosts collected in the 
field and in hosts parasitized in the labo- 
ratory, but in all cases only one larva de- 
veloped per host, hi the laboratory Aride- 
liis nifotestnccus was able to parasitize 2"'\ 
3"-\ and 4* instar host nymphs, as well as 
adults. Younger instars were more suit- 
able for parasitoid development with 95% 
of 3'^' instars parasitized and 85.7% of 2"^^ 
instars parasitized permitting complete 
parasitoid development. The highest mor- 
tality rate recorded for parasitized adults 
was 80.0%. 

Discussion. — Terminology used in the 
description follows that of Sharkey and 
Wharton (1997). This species is a typical 
member of the genus and can be keyed to 
genus without difficulty using the key 
provided by Shaw (1997). This species can 
be identified using the key to Chinese Ar- 
iiiclus species provided by Chen and van 
Achterberg (1997). In the key to world 
species of Papp (1965) this species keys to 
couplet 23, A. iiigritliornx Muesebeck, but 
A. nifotestnccus can be distinguished from 
that species by its lighter colored antenna 
(flagellum entirely black in A. nigrithoriix), 
more coarsely sculptured head (head only 
finely punctate in A. iiigrithonix), and 
weakly developed median frontal carina 
(strongly developed in A. )iigritliomx). A. 
rufotcstnccus can be distinguished from A. 
egrcgrius Schmiedeknecht, the only other 
European species, by its more coarsely 
sculptured head and lighter body (head 
mostly smooth and body black in A. cgrc- 
gius). The three described North America 
species, A. fishcri (Viereck), A. uiclaudcri 
(Brues), and A. iiigrithonix Muesebeck, are 
identical morphologically but differ only 
in color (entirely orange, black head, or 
black mesosoma, respectively). Given the 
wide range of color variations seen in A. 
rufotcstnccus, it would seem questionable 
to separate Aridclus species based on color 
alone. Careful field studies are needed for 
the North American species to examine if 



the observed color forms are related to 
patterns of host use. 

The study of Capek and Davidova-Vi- 
limova (1978) suggested that there are 
four larval instars in A. cgrcgius, but our 
observations suggest only three larval in- 
stars in A. rufotcstnccus. Capek and Davi- 
dova-Vilimova defined their instars 1 and 
2 as morphologically similar, but differing 
only in slight differences in the length of 
the mandible. In fact, since their first instar 
was arbitrarily defined as comprising the 
smallest individuals, and was based only 
on 2 individuals (N = 2), their sample size 
was simply too small to demonstrate a sta- 
tistically significant difference between 
their instars 1 and 2. Another possible ex- 
planation is that all their individuals with 
long, fighter-type mandibles and caudal 
appendage belong to the same instar (1) 
and there are only 3 instars. 

It is worth stressing that in Italy no 
braconid has ever been recorded as para- 
sitoid of Pentatomidae. Moreover, since 
1989, the Department of Arboriculture 
and Plant Protection of the University of 
Perugia periodically collected N. viriduln 
from the field to assess the parasitization 
level of tachinid flies, and the presence of 
any braconid was never observed. In con- 
sideration of this, we can hypothesize a re- 
cent fortuitous introduction of A. iicznrn- 
plingus in Italy, as happened in the past for 
the tachinid Triciiopodn pcniiipcs F. (Colaz- 
za ct nl. 1996). Since the parasitoid was al- 
ready recorded from China and Russia, 
this may be a natural range extension 
from eastern areas. Another possibility is 
that the parasitoid may have previously 
been present but niade a recent host- 
switch from other hosts. However, no al- 
ternate hosts have yet been found in Italy, 
although the following pentatomids have 
been examined for the presence of the par- 
asitoid: Eiirydciun olcrnccum (L.), Eurydciiin 
vcntrnlc (Kit.), Eurygnstcr sp., Grnpliosoiiin 
lincntuiii (L). and G. scmipuiictntuiii (F.). 

MntcrinI cxnniiiicd for rc-dcscriptioii of 
ndiilt. — 3 females: Italy, Palermo, lab 



Volume 10, Number 2, 2001 



137 



reared ex. Nezara viridiila, December 1999; 
5 females, 1 male, same data except Pe- 
rugia, October 1998; 21 females, same data 
except Perugia, August 1999. Specimens 
deposited in University of Wyoming In- 
sect Museum, Laramie; Nationaal Natu- 
urhistorisch Museum, Leiden, The Neth- 
erlands; and Natural History Museum, 
Budapest, Hungary. 

ACKNOWLEDGMENTS 

We are very grateful to Prof. Ferdinando Bin, De- 
partment of Arboriculture and Plant Protection, Uni- 
versity of Perugia, Italy for his helpful cooperation. 
We also wish to thank Dr. C. van Achterberg of the 
Nationaal Natuurhistorisch Museum, Leiden, The 
Netherlands, for his helpful cooperation in borrowing 
the holotype of A. nifotcstacciis and confirming the 
identity of this species. 

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life history of Aridcliis cgrcgiiis (Hymenoptera, 
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J. HYM. RES. 
Vol. 10(2), 2001, pp. 138-144 

<w Species of Streblocera (Asiastreblocera) (Braconidae: 
Euphorinae) from Thailand with Depressed Ovipositor 

Donald L. J. Quicke and Andy Purvis 

(DLJQ) Unit of Parasitoid Systematics, Department of Biology, Imperial College at Silwood 
Park, Ascot, Berkshire SL5 7PY, UK, and Department of Entomology, The Natural History 
Museum, London SW7 5BD, UK; (AP) Department of Biology, hnperial College at Silwood 

Park, Ascot, Berkshire SL5 7PY, UK 



Abstract. — A new species of euphorine braconici, Streblocera (Asiastreblocera) olivera Quicke 
and Purvis, new species, from Thailand is described and illustrated. This is the first record of the 
genus from SE Asia. The description uses SEM to illustrate several features, including the ovi- 
positor and facial horn for the first time. S. [A.) olivera is distinguished from the other four de- 
scribed members of the subgenus. 



Euphorine braconids are interesting be- 
cause they attack adults of holometabo- 
lous insects and adults and nymphs of 
hemimetabolous ones. Probably related to 
this, the adults show a remarkable range 
of forms which are often believed to be 
associated with host manipulation, includ- 
ing remarkably derived antennae and ovi- 
positors (Shaw 1988). 

Streblocera Westwood is a diverse genus 
of principally tropical species that is cur- 
rently divided into a number of subgen- 
era, largely on the basis of the form of the 
antem"ia. The hosts of Streblocera species 
appear to be chrysomelid beetle adults 
(Maeto and Nagai 1985, Shaw 1985), 
though host records are unavailable for 
most species. The subgenus S. (Asiastreblo- 
cera) Belokobylskij was described on the 
basis of a single species from China (S. cor- 
uiita Chao 1964) (Belokobylskij 1987) and 
since then three additional species have 
been described, and collectively the 
known subgeneric range has been found 
to include from Taiwan and Korea (Wang 
1983, Chou 1990, Ku 1997, Belokobylskij 
and Ku 1998). Asiastreblocera differs from 
all other Streblocera in having a facial horn, 
the antenna geniculate at the P' flagellar 



segment which is not serrate but produced 
into a large ventral, flattened lobe, and in 
having a very short ovipositor that is not 
exserted. Recently we were able to study 
a series of a new species of Streblocera 
(Asiastreblocera) from Thailand collected 
by Dr Doug Yanega (University of Cali- 
fornia, Riverside), and this has given us 
the opportunity to obtain DNA sequence 
data for the subgenus, and to study the 
niorphology of its modified antenna and 
ovipositor in more detail using scanning 
electron microscopy. Its D2-D3 28S rDNA 
sequence has been deposited in the EMBL 
database (accession number AJ302831) 
and will be incorporated into a forthcom- 
ing molecular phylogenetic study of the 
non-cyclostome braconids (Belshaw and 
Quicke, in press) and a combined molec- 
ular and morphological phylogeny of the 
Euphorinae (Quicke, Shaw and van Ach- 
terberg in preparation). We are describing 
this species here so as to make its name 
available for future publications. 

TERMINOLOGY AND COLLECTIONS 

Body morphology terminology follows 
Achterberg (1979, 1988); wing venation 
terms used follow Sharkey and Wharton 



Volume 10, Number 1, 2001 



139 



(1997). Collections are abbreviated as fol- 
lows: The Natural History Museum, Lon- 
don (BMNH); University of California, 
Riverside (UCR). 

Streblocera (Asiastreblocera) 
Belokobylskij 

Type species: Streblocera cornuta Chao 1964. 

Diagnosis. — The only euphorine subge- 
nus that has either a facial horn or a single 
or paired ventral projection from the 5"' 
metasomal sternite. See Chen and van 
Achterberg (1997) for additional features 
of the subgenus. 

Distribution. — China, Korea, Taiwan, 
Thailand and Vietnam. 

Streblocera (Asiastreblocera) olivera 
Quicke and Purvis, new species 

(Figs. 1-16) 

Holotype female.— THAILAND: 2km 
south of Ban Pha Bong (a small town 
south of Mae Hong Son), riparian forest, 
low elevation, l.vi.2000, coll. D. Yanega 
(BMNH). Para types. 2 females, both with 
same data as holotype (one coated in plat- 
inum and used for scanning electron mi- 
croscopy BMNH, the other deposited in 
UCR). 

Diagnosis. — This species may be distin- 
guished from all other Streblocera species 
by the possession of a single, non-furcate, 
medio-posterior projection from the 5"' 
metasomal sternite. 

DescriptioiL — Female. Body length 2.6 
mm and of forewing 2.8mm. Antenna ge- 
niculate, 18-segmented, the terminal fla- 
gellomere partially divided on one side; 
scapus 1.56X longer than 1"* flagellar seg- 
ment which is strongly produced ventral- 
ly into lobe; pedicellus with notch ven- 
trally containing discrete row of short, 
erect sensilla (Fig. 7). Scapus with 6-9 di- 
agonal ridges medially (Figs. 2, 6); 1^' fla- 
gellar segment with three diagonal ridges 
on the medio-ventral surface of the pro- 
truding lobe (Fig. 5). Head: 1.8X wider 
than medially long (excluding facial horn); 



1.8x width of mesoscutum; largely 
smoorh. Clypeus distinctly protruding 
from plane of face. Face densely setose 
(Figs. 2, 3). Face as wide as eye height; fa- 
cial horn large, up-curved, apically blunt 
(transverse) in dorsal profile, without 
mid-longitudinal ridge (Figs. 1-3). Cheek 
height 0.26 X height of eye, l.OX basal 
width of mandible in frontal aspect. Dis- 
tance between anterior tentorial pits: 
shortest distance from anterior tentorial 
pit to eye = 2.45:1.0. Width of head 2.35 X 
width of face. Temple strongly narrowed 
behind eyes (Fig. 1). Transverse diameter 
of eye 4.0 X length of temple (dorsal view). 
Eye 1.8X taller than wide in lateral aspect. 
Frons rather flat, medially glabrous, with 
slight pitting behind antennal sockets (Fig. 
1). Transverse diameter of posterior ocel- 
lus: distance between posterior ocelli: 
shortest distance between posterior ocel- 
lus and eye = 1:2:3. Occipital carina ab- 
sent medio-dorsally. Mesosoma: 1.7X lon- 
ger (including neck-like pronotum) than 
maximally high. Pronope well-developed 
(Fig. 4). Mesoscutum smooth and shiny, 
virtually glabrous except along notauli 
which are weakly crenulate anteriorly. 
Notauli deep, largely smooth except for a 
few weak crenulae anteriorly; meeting 
medio-posteriorly in front of scutellar sul- 
cus in a weakly depressed area with a few 
striae but no distinct midlongitudinal ca- 
rina (Fig. 4). Scutellar sulcus long, with a 
single median carina (Fig. 4). Scutellum 
rather convex; medioposteriorly with a 
pair of pits (Fig. 10). Propodeum with me- 
dian carina, on anterior 0.6 and with two 
pairs of strong transverse carinae; postero- 
medially with several short transverse ca- 
rinae (Fig. 10). Fore wing: Vein 1-SR + M 
absent. Vein SRI reaching wing margin 
0.52 of distance from apex of pterostigma 
to the wing tip. Vein SRI 1.9X longer than 
m-cu. Vein 2-CU1 4x longer than 1-CUl. 
Vein r arising 0.63 distance from base of 
pterostigma. Pterostigma 3.0-3.1 X longer 
than wide. Hind xving: Lengths of veins 1- 
M:lr-m:M+CU = 3.0:4.0:1.0. Legs: Length 



140 



Journal of H\ menoptera Research 




Figs. 1-4. Strcbloccm {Asinstn'bloccm) olivera sp. n., female paratype, scanning electron micrographs. \, Head, 
dorsal view. 2, Head, facial view showing also, medioventral aspect of scapus with ridges, and an attached 
insect larva below mandibles. 3, Head, lateral aspect of facial horn. 4, Pronotum and mesoscutum, dorsal 
aspect. Scale bar (see Fig. 3): 1, 4 = 270 (xm; 2 = 430 ixm; 3 = 500 |xm. 



of fore femur: tibia: basitarsus = 2.0:2.35: 
1.0. Hind femur: tibia: basitarsus = 2.2:3.0: 
1.0. Hind tibial spurs almost equal in 
length, each 0.25 X length of hind basitar- 
sus. Mctasoimr. First tergite with fine lon- 
gitudinal striation postero-laterally (Fig. 
11);. remaining tergites completely 
smooth; suture between 2nd and 3rd ter- 



gites almost indistinguishable. 1st tergite 
1.65X longer than maximally wide; maxi- 
mum width 3.1 X minimum width; dis- 
tance from spiracles to posterior margin of 
tergite 1.25X distance between spiracles; 
dorsopes very deep, separated from each 
other by an internal septum, visible 
through cuticle. 2nd tergite with a single 



Volume 10, Number 2, 2001 



141 




Figs. 5-8, Strcbloccm {Asiastrebloccra) olivcra sp. n., female paratype, scanning electron micrographs. 5, 1"' 
flagellar segment showing 3 ridges on dorso-medial surface. 6, Scapus, pedicellus and base of tlagellum, 
arrows indicating ridges. 7, Detail of pedicellus, ventro-lateral aspect showing transverse basal groove with 
row of short sensilla. 8, Anterior mesosoma, lateral aspect. Scale bar (see Fig. 7): 5 = 150 jxm; 6 =^ 270 (xm; 7 
= 30 |jLm; 8 = 380 ixm. 



transverse row of seta subposteriorly (Fig. 
12). Tergites 4 and 5 with distinct medio- 
posterior protuberance (Fig. 12). Sternum 
8 of metasoma with a single, mediopos- 
terior, apically rounded projection. Ovi- 
positor very short, not extending beyond 
the apex of the hypopygium; markedly 



dorso-ventrally depressed, apically trans- 
verse in dorsal profile (Figs. 15, 16). Area 
between hypopygium and terminal ter- 
gites setose (Fig. 16). Colour. Body pale 
brownish yellow, the propodeum medi- 
oposteriorly and the anterolateral parts of 
the 1"' metasomal tergite, somewhat dark- 



142 



Journal of Hymenoptera Research 




Figs. 9-12. Strcblocem (Asiastirhloccrn) oUvcra sp. n., female paratype, scanning electron micrographs. 9, Me- 
sosoma, wings and 1^' metasomal tergite, dorsal aspect. 10, Scutellum, metanotum and propodeum detail. 11, 
1^' metasomal tergite showing fine lateral striations. 12, Metasoma, showing small medio-posterior protuber- 
ances of tergites 4 and 5. Scale bar (see Fig. 11): 9, 11 = 200 |xm; 10, 12 = 100 \i.m. 



ev, the legs and face whitish. Antenna 
brown. 

Mr? /d'.— Unknown. 

Etymology. — Named after Oliver Purvis. 

DISCUSSION 

Most euphorines have moderately to 
highly modified ovipositors that are as- 
sociated with oviposition into adult in- 



sects, and their hosts often have highly 
sclerotized exoskeletons. Thus euphorine 
ovipositors mostly appear adapted to pen- 
etrating intersegniental membranes or 
similar vulnerable places on the host, and 
the typical adaptation is moderate to 
strong lateral compression (see Achter- 
berg and Quicke 2000). A few euphorines 
have solved the problem differently, for 



Volume 10, Number 1, 2001 



143 




Figs. 13-16. Strcbloccrn {Asinstrcbloccra) olivcra sp. n., female paratype, scanning electron micrographs. 13, 
Apex of metasoma, lateral aspect, showing ovipositor (Ov) and protuberance (P) from 5"^ metasomal sternite. 
14, Detail of ovipositor from dorsolateral aspect. 15, Postero-ventral view of apex of metasoma showing 
showing ovipositor (Ov), protuberance (P), and 'secretion' covered seta between ovipositor and ovipositor 
sheaths. 16, Detail of apex of ovipositor showing its dorso-vental compression. Scale bar (see Fig. 15): 13, 16 
= 250 |jLm; 14 = 300 |xm; 15 = 136 [x.m. 



example, species of Spmthicopsis van Ach- 
terberg have a dorso-ventrally depressed 
and apically spatulate ovipositor, though 
unfortunately, their hosts are unknown 
(Chen and van Achterberg 1997). The dor- 
so-ventral compression in some Asiastir- 
blocera, including the new species, is inter- 



esting because it shows that it is possible 
to evolve from lateral compression (as in 
S. (A.) cornufa Chao {vide Chen and van 
Achterberg 1997: Fig. 49)— perhaps 
through an almost cylindrical intermedi- 
ate whose mode of function we can not 
guess at. 



144 



Journal of Hymenciptera Research 



Although there has been speculation 
that the modified antennae in Sircbloccra 
may be involved in holding their beetle 
hosts, there have been no actual observa- 
tions of this. The presence of diagonal stri- 
ations on both the scapus and the ventral 
surface of the 1"' flagellar segment is not 
unique to the new species described here 
(Xuexin X. Chen, personal communica- 
tion) though there function is not known. 
They may be involved in host-restraint/ 
manipulation, but it is also possible that 
they might act as stridules for sound pro- 
duction. 

The setae in the anal area of all speci- 
mens examined are covered with an ap- 
parently congealed substance (Figs. 15, 
16). If this material is a secretory product, 
it be involved in some marking function. 
Unfortunately, such congealed materials 
are often ignored by taxonomists making 
it difficult to assess both their taxonomic 
distribution and how consistent they are 
within a taxon. In addition, S. {A.) olivera 
has the 4"' and 5"' metasomal sternites par- 
ticularly densely setose, but these setae are 
not covered in secretion and their function 
is unknown though they could be sensory. 

ACKNOWLEDGEMENTS 

We are indebted to Doug Yanega (Entomology Re- 
search Museum, University of California, Riverside) 
for collecting this interesting species and making ma- 
terial available for study, to Xuexin X. Chen for in- 
fomation about work on Asinstrcbloccrn in China and 
Korea, to Nicola Simpson for making the SEMs and 
to David Orme for montaging the plates. The speci- 
mens were exported under licence (No. 0741/481) is- 
sued by the Bangkok CITES office of the Royal Forest 
Department. 

LITERATURE CITED 

Achterberg, C. van. 1979. A revision of the subfamily 

Zelinae auct. (Hymenoptera, Braconidae). 

Tidjschrift voor Eniomologie 122: 241-479. 
Achterberg, C. van. 1988. Revision of the subfamily 

Blacinae Foerster (Hymenoptera, Braconidae). 

Zoologischc Vcrliiiiuiclin^cii, Leiden 249: 1-324. 



Achterberg, C. van and D. L. J. Quicke. 2000. The 
palaeotropical species of the tribe Cosmophorini 
Capek (Hymenoptera: Braconidae: Euphorinae) 
with descriptions of twenty-two new species. 
Zoologischc Mcdedcliiigcn, Leiden 74: 283-338. 

Belokobylskij, S. A. 1987. To the knowledge of the 
braconid wasps of the genus Strcblocera Westw. 
(Hymenoptera, Braconidae) of the southern Far 
East. Entoniologisclieskoe Ohozrenie 66: 159-174. 

Belokobylskij, S. A. and D. S. Ku. 1998. Notes on the 
Korean species of the genus Strehloeem with de- 
scriptions of a new species and a key to Korean 
species. The Korean journal of S\/fiteiuatic Zoology 
14:319-325. 

Belshaw, R. and D. L. J. Quicke. In press. Assessing 
character transitions when estimates of phylog- 
eny are uncertain: the evolution of koinobiosis on 
concealed hosts by ichneumonoid parasitoids. 
Systeiuatie Biology. 

Chao, H. 1964. Descriptions of four new species of 
braconid wasps of the genus Strebloeera West- 
wood (Hymenoptera). Aeta Zootaxononiica Siniea 
1: 153-162. (In Chinese with English summary.) 

Chen, X. and C. van Achterberg. 1997. Revision of the 
subfamily Euphorinae (excluding the tribe Me- 
teorini Cresson) (Hymenoptera: Braconidae) 
from China. Zoologisehe Verhandelingen 313: 1- 
217. 

Chou, L.-Y. 1990. The Braconidae (Hymenoptera) of 
Taiwan 11. The genus Strebloeera (Euphorinae). 
lournal of Taiwan Museum 43: 89-148. 

Ku, D. S. 1997. A taxonomic study of the genus Stre- 
bloeera Westwood from Korea. Inseeta Koreana 14: 
65-80. 

Maeto, K. and K. Nagai. 1985. Notes on braconid par- 
asitoids of Medythia nigrobilineata (Motschulsky) 
(Coleoptera, Chrysomelidae), with description of 
a new species of Centistes Haliday (Hymenop- 
tera: Braconidae). Kontyii, Tokyo 53: 729-733. 

Sharkey, M. J. and R. A. Wharton. 1997. Morphology 
and terminology, pp 19-37 in R. A. Wharton, P. 
M. Marsh and M. J. Sharkey (Eds) Manual of the 
New World Genera of the Family Braconidae 
(Hymenoptera). Speeial Pubtieatioii of the Interna- 
tio)\al Society of Hi/iuenopterists, No. 1. 

Shaw, S. R. 1985. A phylogenetic study of the subfam- 
ilies Meteorinae and Euphorinae (Hymenoptera: 
Braconidae). Entoniography 3: 277-370. 

Shaw, S. R. 1988. Euphorine phylogeny: the evolution 
of diversity in host-utilisation by parasitoid 
wasps (Hymenoptera: Braconidae). Eeologieal En- 
tomology 13: 323-335. 

Wang, J.-Y. 1983. A new species of Strebloeera West- 
wood (Hymenoptera: Braconidae: Euphorinae). 
Eiitomotaxonomia 5: 231-232. 



J. HYM. RES. 
Vol. 10(2), 2001, pp. 145-162 

The Afrotropical Species of Leptomastidea Mercet (Hymenoptera: 
Encyrtidae), Parasitoids of Mealybugs 

G. L. Prinsloo 

Biosy sterna tics Division, ARC-Plant Protection Research Institute, Private Bag X 134, 

Pretoria, 0001, South Africa 



Abstract. — The Afrotropical mealybug parasitoids of the genus Lcptoiiiastidca Mercet (Encyrtidae) 
are revised and the following six species described as new: ascia, lamto, poudo, tecta, turba and 
usta. Diagnoses are provided for L. almormis (Girault) and L. jeanneli Mercet, revised status, in 
addition to a key for the separation of females of the eight species of the genus kiiown from the 
region. 



Leptomastidea Mercet is an Old World 
genus, the species of w^hich are all primary 
endoparasitoids of mealybugs (Homop- 
tera: Pseudococcidae). The genus is partic- 
ularly well known through L. abuorniis 
(Girault), which has been used extensively 
in the biological control of the citrus 
mealybug, Planococciis citri (Risso), in sev- 
eral parts of the world, including coun- 
tries in North and South America and Af- 
rica as well as Australia (Noyes and Hayat 
1994). Undoubtedly, species of Leptomas- 
tidea also play a role in regulating mealy- 
bug populations in their native environ- 
ment, as in the case of L. usta sp. nov., an 
indigenous species that is part of the coni- 
plex of hymenopterous parasitoids asso- 
ciated with citrus mealybugs in certain ar- 
eas of South Africa. 

Apart from the six new species de- 
scribed below, Leptomastidea is known 
from 18 species worldwide, the majority 
of which are founci in the Palaearctic and 
Oriental regions. A key to the Palaearctic 
species is provided by Trjapitzin (1989) 
and a detailed account of the Oriental fau- 
na is given by Noyes and Hayat (1994). 
The Afrotrc^pical fauna has not been stud- 
ied in any detail before and was hitherto 
known from only four species: L. almormis 
(Girault), L. jeaiiiieli Mercet, L. sei/rigi Ris- 



bec anci L. aiidKisitreusis Risbec, the latter 
two species having since been transferred 
by Noyes and Hayat (1994) to Homalotylus 
Mayr and Rhitidithorax Ashmead respec- 
tively. Leptomastidea jeanneli was synony- 
mized with L. abnormis by Noyes (2000) 
but is treated here as a valid species. 

In addition to numerous specimens col- 
lected by sweeping and Malaise traps in 
West, East and southern Africa, the pre- 
sent study is based niainly on reared 
South African material. There are at least 
three apparently undescribed species 
among the material that have been exclud- 
ed from this study because of the paucity 
of specimens, whereas the specific identity 
of several other specimens could not be 
determined with certainty. In view of the 
ciifficulties encountered during this study 
in interpreting the nature of certain vari- 
ation it is felt that the identity of these 
specimens is best left in abeyance until ad- 
ditional material, especially host-reared 
series, become available. 

The following acronyms are used in the 
text: BMNH (The Natural History Muse- 
um, London); MNCN (Museo Nacional de 
Ciencias Natu rales, Madrid); NMK (Na- 
tional Museum of Kenya, Nairobi); SANG 
(South African National Collection of In- 
sects, Plant Protection Research Institute, 
Pretoria). 



146 



Journal of Hymenoptera Research 



Leptomastidea Mercet 

Leptomastidca Mercet 1916: 112. Type species 
Leptomastidea aiiraiitiacn Mercet, by monoty- 

py- 

Tanaoiiiastix Timberlake 1918: 362. Type-species 
Pnraleptomastix abnonnis Girault, by original 
designation. 

A detailed account of the taxonomic sta- 
tus of Leptoiiiastiden, including a diagnosis 
and key for separating it from other gen- 
era of the tribe Anagyrini, is provided by 
Noyes and Hayat (1994) and need not be 
repeated here. Suffice it to mention that 
Leptomastidea is most closely allied to Gi/r- 
amisoidea Compere, the eight Afrotropical 
species of which were treated by Prinsloo 
(1983). Noyes and Hayat (1994) state that, 
pending a phylogenetic analysis of the 
Anagyrini, these two genera may eventu- 
ally be considered synonymous. Judging 
by the extent to which certain characters 
overlap between these genera in some ex- 
tra-limital species there may be justifica- 
tion for such a step. However, as far as the 
Afrotropical fauna is concerned, the two 
genera can be readily delineated and 1 
therefore agree with Noyes and Hayat 
(1994) that they should, at least for the 
time being, be treated separately. 

Some of the Afrotropical species treated 
here are structurally very similar, espe- 
cially in taxonomically important charac- 
ters such as the shape and relative dimen- 
sions of the head, antenna, thorax, fore- 
wing venation and ovipositor, and in the 
setation and sculpture of the body. On the 



other hand, these species differ markedly 
in the maculation of the forewing, the ar- 
rangement of fine and coarse setae on the 
wing disc and, in most cases, colour pat- 
tern of the body. The c][uestion is whether 
these differences merely represent infra- 
specific variation and whether those forms 
that are otherwise morphologically very 
similar should therefore be treated as geo- 
graphical races (subspecies) or variants of 
the same species rather than distinct spe- 
cies. This is especially so in the case of L. 
abuoiiuis (Girault) and L. jeauneli Mercet, 
and in the closely allied L. triiba sp. nov. 
and L. iista sp. nov. 

The likelihood of these differences being 
of an infraspecies nature has, however, 
been precluded here. In this regard wing 
maculation and setation in particular were 
found to be stable differentiating charac- 
ters, both within populations and between 
geographically widely separated popula- 
tions, with no evidence of any clinal or 
gradual variation being present. This, cou- 
pled to the fact that the forms in cjuestion 
are evidently sympatric, renders it unlike- 
ly that they are mere variants or races of 
the same species, and they are conse- 
cjuently regarded as representing distinct 
species. This does not imply that all ob- 
served differences in colour and wing 
maculation have been interpreted as being 
of an interspecific nature since infraspecif- 
ic variation, usually in the form of colour 
differences within populations, is evident- 
ly also present, as is commonly found in 
various taxa of the Anagyrini. 



KEY TO AFROTROF^ICAL SPECIES OF LEPTOMASTIDEA 
(based on females) 

1. Forewing hyaline except for an oblique dark cross-band below submarginal vein and a 
dark patch at apex of venation; forewing setation uniform, not forming areas of fine and 
coarse setae (Fig. 1); hind coxa white 8. poiido sp. nov. 

- Forewing maculation different; forewing setation comprising areas of fine and coarse setae 
(Figs. 2-8); hind coxa brown to blackish-brown 2 

2. Forewing with a dark cross-band below submarginal vein and a second, broadly inter- 
rupted band below apex of venation, disc beyond venation hyaline but may appear slightly 



Volume 10, Number 1, 2001 



147 




7. pondo 



5. abnormis 



2. usta 




6. jeannell 





7. turba 



8. ascia 



Figs. 1-8. Lcphvimstiiica spp., female (paratypes except in alvioiDiis and jcanncU), forewing, showing macula- 
tion. 



darkened in part because of the presence of coarse dark setae (Figs. 2, 3); mesopleuron 
orange to red 3 

- Forewing maculation different, wing disc beyond venation partly infuscated (Figs. 4-8); 
mesopleuron always whitish, or pale with dusky suffusions 4 

3. Gena boldly marked with blackish-brown; ovipositor about as long as middle tibia (Fig. 

17) 4. tccta sp. nov. 

- Gena white to yellowish without dark markings; ovipositor about half as long as middle 
tibia (Fig. 13) 3. usta sp. nov. 

4. Apical half of forewing fuscous with a large, oblique hyaline area extending from anterior 
wing margin to near posterior margin as in Fig. 4, extreme apex of wing also hyaline in 
some specimens; c^vipositor about one-third as long as middle tibia (Fig. 26) 

7. lamto sp. nov. 

- Apical half of forewing with maculation different (Figs. 5-8); ovipositor about half as long 

as middle tibia 5 



148 



lOURNAL OF HyMENOPTERA RESEARCH 



5. Forewing with dark cross- band below apex of venation broadly interrupted near posterior 
wing margin as in Figs. 7, 8 6 

- Cross-band below apex of forewing venation complete or narrowly interrupted (Figs. 5, 6) 

7 

6. Thoracic dorsum dark yellow to orange, gaster white basally, blackish apically; forewing 
about 3.0-3.3 X as long as broad, disc beyond venation hyaline with a large subapical dark 
patch at anterior wing margin as in Fig. 7 5. ttirba sp. nov. 

- Thoracic dorsum and gaster entirely blackish-brown to black; forewing broader, less than 
3 X times as long as wide, disc beyond venation with an interrupted subapical cross-band 
band as in Fig. 8 6. ascia sp. nov. 

7. Forewing with dark cross-band below apex of venation at right angles to anterior wing 
margin, parallel-sided; sub-apical cross-band entire, not interrupted in middle (Figs. 6, 9) 

2. jeanneli Mercet 

- Cross-band below apex of venation not parallel-sided but broadening towards posterior 
wing margin; sub-apical cross-band interrupted in middle by a hyaline streak (Fig. 5) .... 

1. ahnormis (Girault) 



1. Leptomastidea abnonnis (Girault) 

(Fig. 5) 

Pamlcptomastix abnonnis Girault 1915: 184. 
Ta)moniastix abnonnis (Girault): Timberlake 

1918: 364. 
Leptomastidea abiionnis (Girault): Mercet 1924: 

255-256. 
Leptomastidea aurantiaca Mercet 1916: 113-116; 

Mercet 1924: 255 (synonymy). 

Diagnosis. — Female. Head yellowish with 
face, temple and gena whitish in most 
specimens; antenna with radicle dark 
brown; scape white below, upper sides 
and dorsal aspect dark brown; pedicel 
with basal half dark brown, apical half 
white; flagellum brownish with basal two 
funicle segments and club usually darker 
than remaining segments. Thoracic dor- 
sum yellowish with mesonotum strongly 
suffused with dark brown; propodeum 
dark brown; side of thorax largely whit- 
ish. Forewing with three dark cross-bands 
as in Fig. 5, intermediate one oblique, 
broadening towards posterior wing mar- 
gin; both intermediate and sub-apical 
bands narrowly interrupted, former near 
posterior wing margin, latter in middle; 
sub-apical band distinctly broader than 
hyaline area separating it from apical 
wing margin. Legs whitish with middle 
and hind coxae brown and dorsal margin 



of all femora darkly outlined; basal half of 
gaster whitish, apical half strongly suf- 
fused with dark brown. 

Male. — Colour, much as in female ex- 
cept: legs entirely white, middle and hind 
coxae not dark; sub-apical cross-band on 
forewing faint, usually indistinct; gaster 
more extensively darkened. 

Material examined. — KENYA: Sabukia, 
xi. 1931, H.C. James, ex Pseiidococcus eitri 
( = prob. Planococciis kemjae (Le Pelley)) on 
Coffea arabica (1 male, det. C. Ferriere). 
SENEGAL: locaHty unknown, v. 1981, J. 
Etienne, ex Ferrisia virgata (Cockerell) (2 fe- 
males, 2 males; T 6324). SOUTH AFRICA: 
Western Cape Province: Paarl, i.l975, G.L. 
Prinsloo, ex Planococciis ficiis (Signoret) on 
grapes (3 females, 1 male; T 5153); Citrus- 
dal, vi.l971, F. Honiball, ex Planococciis ei- 
tri on citrus (2 females, 2 males; T 3854); 
same data except v. 1971 (2 females, 2 
males; T3808); Northern Province: Zebe- 
diela, vi.l966, H. Baas, ex mealybugs on 
citrus (3 females, 1 male; T 2342); Mpu- 
malanga Province: Nelspruit, v. 1972, H.P. 
Insley, with scale insects on Maytenus sp. 
(1 female, 1 male; T 4353); North West 
Province: Rustenburg, xi.l971, C.J. Cilliers, 
ex mealybugs on citrus (3 females, 1 male; 
T 4014); same data except i.l972 (5 fe- 
males, 1 male; T 7n0); Gauteng Province: 



Volume 10, Number 2, 2001 



149 




Figs. 9-13. Lcptomaslidca spp., females. 9, L. jcaiiiicli, lectotype, forevving, showing maculation. 10-13. L. iistn, 
paratypes. 10, Head, frontal view. 11, Mesonotum with sculpture of mesoscutum and scutellum enlarged. 12, 
Apex of forewing venation. 13, Ovipositor and middle tibia drawn to the same scale. 



Pretoria, vi.l965, S.W. Broodryk, host un- 
known (1 female, 1 male; T 1980). UGAN- 
DA: Kiki, vii.1971, K. Ogwaro, with scale 
insects on citrus (1 female; T3912). All 
specimens in SANC. 

Extra-Umital M^ift'rm/.— AUSTRALIA: 
Palmwoods, Queensland, iii.1978, D. Mur- 
ray, ex Planococcus citri on custard apple 
(6 females; T 5269). ISRAEL: Neot Haki- 



kar, xi.l980, Y. Ben-Dov, with scale insects 
on Phoenix dactylifera (2 females, 1 male; 
T6232). USA: "Whittier, Calif. 1922, Rust, 
ex Pscudococcus citri" (10 females, 10 
males; T 4274). All specimens in SANC. 

RcDuirks. — This well known economical- 
ly important species was originally de- 
scribed from Sicily and subsequently in- 
troduced to various parts of the world (in 



150 



Journal of Hymenoptera Research 



many cases via laboratory stocks obtained 
from California) for the control of Piano- 
coccus citri and other mealybug species. A 
summary of the literature pertaining to 
the use of L. nlvioniiis in biological control 
worldwide is provided by Noyes and 
Hayat (1994). 

Within the African context, L. nbiionuis 
was imported into South Africa between 
1934 and 1940 for the control of Phviococ- 
ciis ficus (thought to be P.citri) on grapes 
in the Western Cape Province where it be- 
came established. It was never cieliberate- 
ly released against P. citri on citrus but is 
now found in association with this pest 
throughout the citrus-growing areas of the 
country. Other introductions into Africa, 
which took place during the first half of 
the previous century and regarded as hav- 
ing been unsuccessful (see Noyes and 
Hayat 1994), were to Ghana and Kenya for 
the control of Plniiococcoidcs njnlcnsis 
(Laing) and Plniiococcus keiiyac respective- 

ly- 

Lcptomasiidca nbnoruiis has repeatedly 
been redescribed and illustrated in the lit- 
erature, with recent accounts by Noyes 
(1988, 2000). Suffice it to mention that this 
species can be distinguished from its Af- 
rican congeners by the foregoing diagno- 
sis and key. Its relationship with L. jcniincli 
Mercet, with which it was synonymyzed 
by Noyes (2000), is discussed in the treat- 
ment of the latter species below. 

2. Leptoniastidea jeanucli Mercet, 
revised status 

(Figs. 6, 9) 

Lcptoiiiastidcii jcaniicli Mercet 1924: 256-258; 
Compere 1939: 25; Noyes 2000: 138 [as a ju- 
nior synonym of L. abnormis (Girault)]. 

RcdescriptioiL — Card-iuoiiiitcd female lec- 
totypc. Length: 1.0 mm. Colour. Head en- 
tirely whitish, obviously faded; antenna 
uniformly blackish-brown except apical 
one-third or so of pedicel white. Thorax, 
propodeum and gaster blackish-brown ex- 
cept mesopleuron and prepectus whitish. 



tegula white basally, darkly suffused api- 
cally. Forewing with three dark cross- 
bands as in Fig. 9, intermediate one at 
right angles to anterior wing margin, rel- 
atively narrow, parallel-sided, not some- 
what oblique and broader towards poste- 
rior wing margin as in L. abnoriuis; inter- 
mediate and sub-apical bands not inter- 
rupted; sub-apical band subequal in width 
to hyaline area separating it from apical 
wing margin. Legs with fore coxa whitish, 
middle and hind coxae blackish-brown; 
legs otherwise whitish with hind femur 
entirely blackish-brown and fore and mid- 
dle femora darkly outlined dorsally. Head: 
in dorsal view, twice as wide as frontov- 
ertex at median ocellus; ocelli in a right- 
angled triangle, lateral pair about twice 
own diameter from eye margins; head 
otherwise typical of genus, inner eye mar- 
gins approximately parallel-sided, fron- 
tovertex with regular, raised reticulate 
sculpture; eyes appearing naked. Antenna 
with scape subcylindrical, about 5.5 X as 
long as wide; pedicel as long as basal fu- 
nicle segment; funicle segments subequal 
in length, basal one 3.5 X as long as wide, 
remaining segments becoming progres- 
sively very slightly broader; club slightly 
longer than distal two funicle segments 
combined; linear sensillae discernible on 
funicle segment VI and club. Forewing 
2.9 X as long as wide; costal cell hardly 
discernible; basal triangle of wing disc 
densely and fairly coarsely setose except 
for a narrow bare streak below basal third 
of submarginal vein, this streak separated 
from submarginal vein by a single row of 
setae; areas of wing disc delineated by fus- 
cous cross-bands with dark, fairly coarse 
setae, setae covering hyaline areas fine 
and pale, hardly visible under low mag- 
nification; postmarginal vein approxi- 
mately 3x as long as marginal, a little less 
than twice as long as stigmal vein. Middle 
leg with tibial spur a little shorter than 
basal tarsal segment. General shape of 
thorax and gaster similar to A. abnoriuis 
and many other species of the genus. 



Volume 10, Number 1, 2001 



151 



sculpture and setation of thorax not clear- 
ly discernible in the card-mounted speci- 
men, but appearing similar to that of the 
latter species. 

Variation. — Female. At hand are a num- 
ber c^f specimens from Zimbabwe, Kenya 
and various countries in West Africa 
which differ in colour front the female lec- 
totype as follows: antenna with scape and 
flagellum not uniformly dark but scape bi- 
colorous, whitish below, upper sides and 
dorsum blackish-brown, funicle entirely 
pale brown or with basal two or three seg- 
ments distinctly darker, club blackish- 
brown; thoracic dorsum dominantly yel- 
lowish-brown; basal half or so of gaster 
white, apical half blackish-brown; legs, 
save dark middle and hind coxae, entirely 
whitish. Also available is a single addi- 
tional specimen from Zimbabwe which, 
unlike these speciniens, differs from the 
lectotype only in the paler thoracic dor- 
sum and bicolorous antennal scape. 

Type material exaiuiiied. — Female lecto- 
type (MNCN), designated by J.S. Noyes, 
with following data: "Naivasha, Africa or. 
Inglesa; Leptomastidea jeanneli Mercet, 
tipo; MNCN Cat. Tipo No. 10434". 

Additional material. — GHANA: Tafo, iv 
and V. 1973, M. Bigger, ex Plaiioecoides iija- 
leusis (9 females BMNH). KENYA: "Don- 
yo, Sabuk, 1939, C 127, A.R. Melville, ex 
Pseudococcus sp. on Combretum, B.M. 
1839-601, Leptomastidea jeanneli Mercet, 
det. Ferriere" (4 females; T 6621; SANC). 
NIGERIA: Ibadan, IIT Compound, xi.l987, 
J.S. Noyes (19 females; BMNH). TOGO: 5 
km. W Amiame, 16.xii.l988, J.S. Noyes (1 
female; BMNH). SAO TOME: xii.1974, J.O. 
Derron, ex Plauoccoides iijaleiisis on cacac^ 
(1 female; T 4943; SANC). ZIMBABWE: 
Harare ( = Rhodesia: Salisbury), xi.l974, 
ix.l976, ix.l979, iii.1984, A. Watsham (4 fe- 
males; BMNH). 

Remarks. — This species was originally 
described from an undisclosed number of 
female specimens from Naivasha, which is 
in Kenya, not Uganda as cited by Mercet 
(1924). The only known type specimen, a 



lectotype designated by Noyes (1981), is in 
good condition and mounted on a card. 

Leptomastidea jeanneli was recently syn- 
onymyzed with L. abnormis by Noyes 
(2000) who mentioned that the characters 
listed by Mercet (1924) for separating 
these two species fall within the range of 
variation found in L. abnormis. Although 
these two species are indeed very similar, 
I am nevertheless of the opinion that L. 
jeanneli should be surrected as a valid spe- 
cies. I base this decision on the distinct dif- 
ference in the shape of the fuscous cross- 
bands on the forewing, as described above 
and shown in Figs. 5, 6 and 9. I do not 
believe that this difference reflects infra- 
specific variation, especially in view of the 
fact that the characteristic wing pattern in 
L. abnormis is not known to vary signifi- 
cantly, as is evident from the large amount 
of available study material and many pub- 
lished accounts of this geographically 
widespread and experimentally well 
known species. 

The difference in body colour between 
the female lectotype of L. jeaivieli and the 
specimens mentioned under "Variation" 
is attributed to infraspecific variation on 
the basis of the intermediate colour pat- 
tern of the single specimen from Zimbab- 
we. In this specimen the gaster, legs, and 
antennal flagellum are characteristic of the 
lectotype, whereas the antennal scape is 
bicolorous as in the other specimens from 
Zimbabwe and those from West Africa 
and Kenya. I am also of the opinion that, 
as far as the noted variation in colour is 
concerned, both the lectotype and inter- 
mediate specimen from Zimbabwe repre- 
sent aberrant forms of L. jeanneli, the "nor- 
mal" form being represented by the re- 
maining specimens. Intel estingly, the se- 
ries from Kenya was identified as L. 
jeanneli by the late Ch. Ferriere, lending 
further support to the present interpreta- 
tion of the taxonomic identity of the study 
material. 

Compere (1939) recorded this species 
from a short series of specimens from 



152 



Journal of Hymenoptera Research 



Kenya and a single female from Eritrea, 
none of which, according to Compere, are 
in complete agreement with Mercet's de- 
scription of L. jeanucU. I have not seen 
these specimens. 

3. Leptomastidea usta Prinsloo, new 
species 

(Figs. 2, 10-15) 

Description. — Fcinnlc. Length: 0.7-0.9 
mm. Colour: Head, thorax and propodeum 
entirely reddish-orange to red except teni- 
ple, gena, n^iouth margin and area be- 
tween scrobes white to yellowish; tegula 
white to reddish basally, brown apically; 
setae on frontovertex and mesonotum 
white. Antenna with radicle blackish- 
brown; scape white below, upper sides, 
and dorsal aspect from base to near apex, 
blackish-brown; basal half of pedicel 
blackish-brown, apical half white; flagel- 
lum usually brown to blackish-brown ex- 
cept funicle segments Ill-V or III-VI white; 
tlagellum rarely entirely dark. Forewing 
(Fig. 2) with two dark cross-bands, basal 
one (below subniarginal vein) strongly 
oblique, distal one (at apex of venation) 
broadly interrupted, wing disc beyond ve- 
nation hyaline but appearing slightly 
darkened in part because of the presence 
of coarse dark setae; hind wing entirely 
hyaline. Legs with fore coxa whitish, mid- 
dle and hind coxae blackish-brown; legs 
otherwise mostly whitish with dorsal mar- 
gins of all femora darkly outlined, middle 
and hind tibiae also somewhat darkened 
in some specimens. Gaster white with api- 
cal third unevenly suffused with brown to 
blackish-brown, base also dark in some 
specimens. Head: in dorsal view, 1.7-1.9X 
as wide as frontovertex at median ocellus; 
ocelli in a right-angled triangle, lateral 
pair separated from inner eye margins by 
about twice own diameter; in frontal view 
(Fig. 10), about 1.2X as wide as long, ma- 
lar space 0.5 X eye length; with regular 
more or less circular sculptural cells as il- 
lustrated, the diameter of cells on frontox- 
ertex about the same as that of eye facet; 



front aspect of head, and eyes, sparsely 
and finely setose, eyes appearing naked 
under low magnification, setae shorter 
than diameter of eye facet. Antemia (Fig. 
15) with scape subcylindrincal, approxi- 
mately 5x as long as wide; pedicel rang- 
ing from as long as to 1.3X as long as bas- 
al funicle segnient; funicle segments sub- 
equal in size, basal segment 2.5-3.1 X as 
long as wide; club as long as distal two 
and a half funicle segments combined; lin- 
ear sensillae on all club and distal two or 
three funicle segn^ients. Thorax: typical of 
genus, dimensions, sculpture and setation 
of mesonotum as in Fig. 11. Forewing (Fig. 
2) 2.8-3.3 X as long as wide; costal cell nar- 
row, barely discernible in basal half in 
most specimens, with a single row of setae 
on ventral margin; venation (Fig. 12) with 
postmarginal vein 1.3-1.6X length of stig- 
mal, 2.5-3.0 X as long as marginal, latter 
0.5-0.6 X length of stigmal vein; setae in 
area between the two fuscous bands and 
in a large patch beyond venation paler 
and much finer than remaining discal se- 
tae. Ovipositor (Fig. 13): 0.5-0.6 x length of 
middle tibia; gonostyli 0.5-0.7X length of 
middle tibial spur, latter subequal in 
length to basal tarsal segnient of middle 
leg. 

Male. — Colour: Head with frontovertex 
c^range, fading to yellow on face and gena; 
antenna with scape whitish except dorsal 
aspect brown; remainder of antenna 
brown. Thorax, propodeum and gaster 
dominantly dark brown to blackish- 
brown with yellowish-brown suffusions 
on sides of n^iesonotum. Forewing with an 
oblique pale brown band below submar- 
ginal vein as in female and a faint dark 
patch at apex of venation that does not 
form a cross-band. Legs much as in fe- 
male. Structure: Differing from female 
mainly as follows: torulus placed higher 
on face, its lower margin more or less level 
with lower eye margins. Antenna (Fig. 14) 
with scape about 4.5 x as long as wide; fu- 
nicle segments subequal in length, each 
about 4X as long as wide; club unseg- 



Volume 10, Number 2, 2001 



153 




Figs. 14-18. Leptomastidea spp. 14-15. L. usta, paratypes. 14, Antenna and sub-basal row of spine-like setae 
on club, male. 15, Antenna, female. 16-18. L. tecta, female paratype. 16, Apex of forewing venation. 17, Ovi- 
positor and middle tibia drawn to the same scale. 18, Antenna. 



merited, as long as distal two funicle seg- 
ments combined; funicle with long, 
curved setae, each about 3x as long as the 
width of a segment; club, apart from nor- 
mal setatic)n, with a longitudinal row of 3- 
4 short straight setae ventrally near base 
(Fig. 14). Forewing slightly broader than 



in female, about 2.5 X as long as wide; 
wing disc uniformly setose, not differen- 
tiated into areas with fine and coarse setae 
as in female. Phallobcise less than half as 
long as middle tibia, digiti each terminat- 
ing in two short, stout booklets. 

Material examined. — Female holotype, 29 



154 



Journal of Hymenoptera Research 



female, 12 male paratypes as follows: 
BOTSWANA: Serowe, ix.l987, P. Forsch- 
hammer (1 female; BMNH). SOUTH AF- 
RICA: Northern Province: Zebediela, vi- 
vii.1966, H. Baas, with Nipmccoccus viridis 
(Newstead) and Piirncoccus bunicnic 
(Brain) on citrus (holotype, 5 females, 1 
male; T 2343); Zebediela, vi.l981, M. van 
der Kooij, ex mealybugs on citrus (6 fe- 
males, T 6307); Western Cape Province: 
De Dooms, 8.x. 1968, V.B. Whitehead, ex 
mealybugs on Elytropappus rliiiioccrotis 
(L.f.) Less. (1 female, 6 males; T 2763); 
same data except H.P. Insley, 14. ii. 1969 (1 
female, 2 males; T 2976); Stellenbosch, 
ii.l969, H.P. Insley, ex mealybugs on Stoc- 
be vulgaris Levyns (2 females; T2975); Stel- 
lenbosch, ix.l965, W.B. Whitehead, ex 
Plioiacoccus stclli (Brain) on Lcucndoniroii 
daphnoidcs (Thunb.) Meisn. (1 female; T 
2019); Nature's Valley, iii.1970, H.P. In- 
sley, ex mealybugs on Metalasia uiuricatn 
(L.) D.Don (2 females; T 3407); Gauteng 
Province: Roodeplaat Dam, nr Pretoria, 
iii.1972, H.P. Insley, ex Delottococcus quac- 
situs (Brain) on Acncia sp. (2 females; T 
4293); Kwazulu-Natal: Oribi Gorge, i.l972, 
H.P. Insley, ex mealybugs on Cryptocan/n 
weyliei Stapf (4 females, 1 male; T 4191); 
Eastern Cape Province: Willowmore, 
i.l979, C. Kok, with Tiiclinrdiiin sp. on £/i/- 
tropappus rJiiuoccrotis (L. f) Less. (4 females, 
2 males; T 7 111). ZIMBABWE: Harare ( = 
Rliodesia; Salisbury), xi.l976, A. Watsham 
(1 female; BMNH). All specimens in 
SANC unless otherwise noted. 

Remarks. — This widespread southern 
African species is readily separated in the 
female from its Afrotropical congeners, 
except L. tccta, by the striking reddish 
head and thorax and maculation of the 
forewing; it differs from L. tccta sp. nov. 
as mentioned in the treatment of that spe- 
cies below. Lcptoinnstidca iista also resem- 
bles L. rubra Tachikawa, which is known 
from the Palaearctic region, closely in 
structure, body colour and general colour 
pattern of the forewing. The two species 
can, however, be distinguished by the 



maculation of the forewing as follows: in 
L. rubra the infuscation at the apex of the 
venation forms a strongly oblique, incom- 
plete band that extends about half-way 
across wing disc; in L. usta this band is 
almost at right angles to the anterior wing 
margin and, although broadly interrupt- 
ed, extends across the entire width of the 
wing. 

4. Leptoutastidea tecta Prinsloo, new 
species 

(Figs. 3, 16-18) 

Description. — Female. Length: 0.7-0.8 
mm. Colour: Head brownish-yellow to 
pale orange with temple white, gena bold- 
ly marked with blackish-brown, this col- 
our extending upwards onto hind margin 
of temple; setae on front aspect of head 
whitish. Antenna with radicle blackish- 
brown; scape with dorsal aspect and up- 
per half of sides blackish-brown, other- 
wise white; pedicel with basal two-thirds 
blackish-brown, fading to white distally; 
flagellum entirely and uniformly blackish- 
brown. Thorax and propodeum orange 
except prepectus and collar of pronotum 
whitish, anterior n^argin of mesoscutum 
(partly obscured by pronotum) darkly suf- 
fused; tegula whitish to orange basally, 
dusky distally; thoracic setation silvery- 
white. Forewing (Fig. 3) with an oblique 
dark band below submarginal vein and a 
second, broadly interrupted one at apex of 
venation, disc beyond venation hyaline, 
appearing partly somewhat darkened be- 
cause of the presence of coarse dark setae; 
hind wing entirely hyaline. Fore coxa 
niostly whitish, middle and hind coxae 
brown; fore leg white with dorsal and 
ventral margins of femur darkly outlined, 
tibia slightly embrowned in one specimen; 
middle and hind legs whitish with dorsal 
margin of femora darkly outlined and 
hind tibia with dusky suffusions, distal 
one or two tarsal segnients dark. Gaster 
white with apical one-third or so blackish- 
brown, extreme base also darkly suffused 
in some specimens. Head: 1.8X as wide as 



Volume 10, Number 2, 2001 



155 



frontovertex at median ocellus, dimen- 
sions, sculpture and setation much as de- 
scribed and illustrated for L. ustn. Antenna 
(Fig. 18) with scape 5.5 X as long as broad; 
pedicel slightly longer than basal funicle 
segment; funicle segments subequal in 
length, basal segment 3.0-3.3 x as long as 
wide; club as long as distal two and a half 
funicle segments combined; flagellum 
finely and fairly sparsely setose as in Fig. 
18; linear sensillae present on club and 
distal three funicle segments in two slide- 
mounted specimens. Tliornx: sculpture 
and setation not significantly ciifferent 
from that of L. ustn. Forewing as in Fig. 3, 
3x times as long as wide, arrangement of 
fine and coarse setae much as in L. ustn; 
costal cell narrow but clearly discernible; 
venation (Fig. 16) with postmarginal vein 
1.3-1.4X as long as stigmal, about 3.2 x as 
long as marginal, latter 0.5 X length of 
stigmal. Ovipositor (Fig. 17): unusually 
long, about as long as gaster and equal in 
length to middle tibia, gonostyli 1.0-1.2X 
as long as middle tibial spur. 

Mnlc. — Colour: Head with frontovertex 
dark yellow, fronto-occipital margin nar- 
rowly outlined in black; face and temple 
white, gena with a bold blackish-brown 
marking as in female. Antenna with scape 
bicolorous as in female, otherwise entirely 
brown. Thoracic dorsum dominantly 
blackish-brown except collar of pronotum 
white, side of mesoscutum and postero- 
lateral margins of scutellum narrowly out- 
lined in orange, metanotum orange; side 
of thorax with mesopleuron boldly 
marked with yellow or orange anteriorly, 
otherwise blackish-brown; prepectus 
white. Forewing hyaline with a pale 
brown oblique band below submarginal 
vein as in female and a large infuscated 
patch at apex of venation. Legs much as 
in female. Gaster entirely blackish. Struc- 
ture: siniilar to male of L. ustn except phal- 
lobase relatively longer, a little more than 
half as long as middle tibia. 

Material examined. — Female holotype, 3 
female, 4 male paratypes (SANC) as fol- 



lows: SOUTH AFRICA: Eastern Cape 
Province: Georgida, xi. 1983, G.L. Prins- 
loo, ex Lennnia sp. on Salsola glabrescens 
Burtt Davy (T 7112). 

Remarks. — This species, which is known 
only from South Africa, can be separated 
in the female from all the other Afrotrop- 
ical species by the unusually long ovipos- 
itor, whereas the forewing maculation is 
unlike that of any of the extra-limital spe- 
cies. Leptomastidea tecta resembles L. usta 
closely in structure, colour and, perhaps 
most significantly, maculation and seta- 
tion of the forewing. It differs from the lat- 
ter species in the genae which are boldly 
marked with blackish-brown, the dimen- 
sions of the antennal segments, and ovi- 
positor which is as long as the middle tib- 
ia, not half as long; correspondingly, the 
phallobase of the male of L. tecta is longer 
than in L. usta. 

5. Leptomastidea turba Prinsloo, new 
species 

(Figs. 7, 19-21) 

Description. — Female. Length: 0.8-1.1 
mm. Colour: Head with frontovertex dark 
yellow, fading to white on lower face, 
temple and gena, fronto-occipital margin 
very narrowly darkened; setae silvery- 
white. Antenna with radicle blackish- 
brown; scape with dorsal aspect and up- 
per sides blackish-brown, otherwise 
white; pedicel blackish-brown basally, 
apical half white; flagellum either entirely 
dark brown to blackish-brown or with fu- 
nicle segments III and IV slightly to dis- 
tinctly paler than remaining flagellar seg- 
ments, this variation present in same sam- 
ple of reared specimens. Thoracic dorsum 
and propodeum ranging from dark yellow 
to orange, pronotal collar white, sides of 
propodeum behind spiracles blackish- 
brown; tegula largely whitish, only apex 
dark; mesonotal setation silvery-white; 
side of thorax whitish. Forewing (Fig. 7) 
with an oblique fuscous cross-band below 
submarginal vein, a second, broadly inter- 
rupted one below apex of venation and a 



156 



Journal of Hymenoptera Research 




Figs. 19-24. Lcptoiiiastidca spp., female paratypes. 19-21. L. tiirhn. 19, Apex of forewing venation. 20, Scutel- 
lum with sculpture enlarged. 21, Antenna. 22-24. L. nficin. 22, Apex of forewing venation. 23, Scutellum with 
sculpture enlarged. 24, Antenna. 



large infuscated patch at anterior margin 
near apex of wing, setae below this infus- 
cated patch darkened, but not wing mem- 
brane; hind wing hyaline. Legs whitish ex- 
cept middle and hind coxae, dorsal mar- 
gins of femora usually, and tarsal tips. 



brown to blackish-brown; gaster with bas- 
al half or so white, apical half blackish. 
Head: 1.9-2.1 X as wide as frontovertex at 
median ocellus; lateral ocelli about twice 
own diameter from inner eye margins; eye 
a little more than twice as long as malar 



Volume 10, Number 2, 2001 



157 



space; sculpture much as described and il- 
lustrated for L. iistn, diameter of cells on 
frontovertex about equal to that of eye fac- 
et; frontovertex from median ocellus to oc- 
cipital margin with numerous scattered 
setae, remainder of front aspect of head 
sparsely setose; eyes finely and inconspic- 
uously setose, setae shorter than diameter 
of eye facet. Antenna (Fig. 21) with scape 
4.7-5.0 X as long as broad; pedicel sube- 
qual in length to basal funicle segment; fu- 
nicle segments subequal in length, basal 
one 3.0-3.5 X as long as wide; club slightly 
longer than distal two funicle segments 
combined; funicle usually with linear sen- 
sillae on all except basal one or two seg- 
ments, rarely present on all segments. T/;o- 
;v?.y: dimensions, sculpture and setation of 
mesonotum much as in L. iistn, cells on 
anterior part of scutellum irregular in 
shape, as shown in Fig. 20. Forewing (Fig. 
7) about 3x as long as wide; costal cell 
very narrow, indiscernible in some speci- 
mens; venation (Fig. 19) with postmargin- 
al vein 1.3-1.4X as long as stigmal, about 
3.5 X length of niarginal vein, latter 0.5 X 
as long as stigmal; wing disc densely and 
evenly setose, arrangement of fine and 
coarse setae as illustrated. Ovipositor: al- 
most 0.5 X length of middle tibia, gonos- 
tyli 0.5 X as long as middle tibial spur. 

Male. — Colour: Differing from female 
mainly as follows: mesonotum not uni- 
formly dark yellow but suffused with 
blackish-brown to a varying degree, gaster 
entirely blackish, its basal half not white; 
forewing with infuscation below apex of 
venation not forming a well defined inter- 
rupted cross-band, but appearing as two 
faint patches, one directly below venation, 
the other at posterior wing margin; disc 
beyond venation hyaline, without a sub- 
apical infuscated patch. Similar to male of 
L. iistn in colour and structure, differing 
only in presence of a faint fuscous patch 
at posterior margin of forewing. 

Variation. — At hand are several female 
specimens collected by sweeping from the 
same locality (Harare) in Zimbabwe. 



These specimens have been excluded from 
the type material, from which they differ 
as follows: front aspect of head entirely 
yellow, not fading to white on face and 
gena; funicle segments IIl-V white in con- 
trast to remaining flagellar segments, 
which are blackish-brown; side of thorax 
not entirely white, mesopleuron with 
dusky suffusions; legs with femora entire- 
ly pale, dorsal margins not outlined in 
blackish-brown; scutellum anteriorly with 
sculptural cells more regular in shape than 
shown in Fig. 20; forewing narrower, 
about 3.3 X as long as wide. 

These differences are here attributed to 
infraspecific variation, although further 
material, including reared series of both 
sexes, is required to determine the exact 
nature of the variation seemingly present 
in L. tiirba. This is especially important in 
view of the presence of a further specimen 
from Harare which, unlike the above- 
mentioned material from this locality, 
does not differ from the type specimens. 

Type material examined. — Female holo- 
type, 36 female, 13 male paratypes as fol- 
lows: NAMIBIA: Otavi, ii.l978, C. Kok ex 
mealybugs (10 females, 1 male; T 7114); 
Chorixas, ii.l978, C. Kok ex mealybugs on 
Welcuitchia mirabilis Hook. f. (3 females; T 
6061). SOUTH AFRICA: Gauteng Prov- 
ince: Pretoria, iv.l995, O.C. Neser, ex Par- 
acoeeiis burnerae on Seneeio venosus Harv. 
(holotype, 16 females, 11 males; T 7113); 
same data except xi.l988, S. Neser (3 fe- 
males, 1 male; T 6978). ZAMBIA: 15 KM 
e Lusaka, 11-19 and 20-31. ii. 1980, 
R.A.Beaver (3 females; BMNH). ZIMBAB- 
WE: Harare, vii.1982, A. Watsham (1 fe- 
male; BMNH). Holotype and paratypes in 
SANC unless otherwise noted. 

Non-ti/p}e material. — ZIMBABWE: Harare 
( = Rhodesia: Salisbury), xii.J978, i-iii and 
viii.1979, xi.l980, A. Watsham, by sweep- 
ing (23 females; BMNH). 

Remarks. — This widespread southern 
African species can be separated, in the fe- 
male, from all other species of the genus 
by the maculation and setation of the fore- 



158 



Journal of Hymenoptera Research 



wing. Structurally it is very similar to L. 
listn, from which it seems to differs only 
in the presence of linear sensillae on one 
or more of the basal three funicle seg- 
ments. Apart from the difference in wing 
maculation, the female of L. tiirbn also dif- 
fers from L. ustn in the colour of the head 
and thoracic dorsum which are paler, and 
the mesopleuron which is white, or white 
with dusky suffusions, not reddish. 

6. Leptojiiastidca ascia Prinsloo, new 
species 

(Figs. 8, 22-24) 

Description. — Female. Length: 1.1-1.4 
mm Colour. Frontovertex orange, fronto- 
occiptal margin broadly suffused with 
blackish-brown, remainder of front aspect 
of head white with gena slightly dark- 
ened; setae on frontovertex and eyes dark. 
Antenna uniformly blackish-brown except 
ventral aspect and lower sides of scape, 
and apex of pedicel, white. Thorax and 
propodeum blackish-brown to almost 
black except: pronotal collar and prepec- 
tus white, tegula whitish basally, other- 
wise dark, mesopleuron entirely whitish, 
or with dusky suffusions; mesonotal setae 
silvery-white. Forewing with three fus- 
cous cross-bands as in Fig. 8, basal one 
oblique, intermediate one (at apex of ve- 
nation) at right angles to anterior wing 
margin, parallel-sided, broadly interrupt- 
ed, the sub-apical band with two narrow 
interruptions; hind wing hyaline with a 
narrow oblique brownish cross-band near 
base. Fore coxa white, middle and hind 
coxae blackish-brown; legs otherwise usu- 
ally sordid white with dorsal margins of 
all femora dark; in some specimens the 
femora, tibiae, and tarsi are more exten- 
sively darkened. Gaster entirely blackish- 
brown. Henii: about twice as wide as fron- 
tovertex at median ocellus; lateral ocellus 
1.5-2. Ox its diameter from lateral eye mar- 
gin; head with shape and sculpture oth- 
erwise much as illustrated for L. iistii ; 
frontovertex fairly densely and strongly 
setose; eye densely and strongly setose, se- 



tae readily discernible under low magni- 
fication, each seta about as long as the di- 
ameter of eye facet. Antenna (Fig. 24) with 
scape just more than 6x as long as wide; 
pedicel subecjual in length to basal funicle 
segment; funicle segments subequal in 
length, basal segment about 3.5 X as long 
as wide; club as long as distal two funicle 
segments combined; funicle segments III- 
VI and club with linear sensillae. Thorax: 
shape of sculptural cells on anterior part 
of scutelluni as in Fig. 23, more regular in 
shape than in the other species treated 
here. Forewing (Fig. 8) 2.7-2.8 X as long as 
broad; costal cell relatively broad, clearly 
visible along its entire length with a single 
row of setae ventrally; venation (Fig. 22) 
with postmarginal vein 1.6 X as long as 
stigmal, 3x as long as marginal, latter 
0.5 X as long as stigmal; setae (save those 
at base) confined to hyaline areas of disc 
finer and paler than those of infuscated ar- 
eas. Ovipositor: 0.5 X length of middle tibia; 
gonostyli 0.6 X as long as middle tibial 
spur, latter subequal in length to basal tar- 
sal segment of middle leg. 

Male. — Colour: as in female except fron- 
tovertex more extensively suffused with 
blackish-brown, and infuscated areas on 
forewing paler, sub-apical cross-band in- 
distinctly delineated. Structure: Differing 
from female mainly by slightly broader 
forewing, which is about 2.5 X as long as 
broad, and antenna; antenna similar to 
that of male of L. usta, as shown in Fig 14; 
digiti of phallobase each terminating in 
two short, stout booklets. 

Matertial examined. — Female holotype, 
55 female, 17 male para types as follows: 
KENYA: "Aberdare NP, 0.23S 36.46E, 10- 
18. ii. 1999, T.Wagner, Canopy fog Podocar- 
pus latifolius, BMNH 1999-279"(holotype, 
27 females, 7 males); "Gatamayu, Kikuyu 
Esc. 2320m, 0.58S 36.42E, T.Wagner, ii.99. 
Canopy fog Podocarpus latifolius, BMNH 
(E) 1999-279" (26 females 7 males); 
"1600m, Mt Kenya NP (WHQ), O.IOS 
37.10E, ii.l999, T, Wagner, Canopy fog Po- 
docarpus latifolius, BMNH (E) 1999-279" 



Volume 10, Number 2, 2001 



159 



(2 females, 3 males). Holotype and para- 
types in NMK; paratypes in BMNH and 
SANC. 

Remarks. — The three series on which 
this new species is based are all from cen- 
tral Kenya where they were collected (by 
fogging) on Podocarpnis Intifoliiis (Thunb.) 
R.Br, ex Mirb., a gymnosperm which is 
widespread in southern and East Africa. 
The mealybug host of this material is un- 
known, but may be Enstin joiibcrti De Lot- 
to, the only species of mealybug known to 
be associated with this tree in sub-Saharan 
Africa. 

Leptouiastidea ascin appears to be most 
closely allied to L. iistn and L. tiirba and, 
as in these two species, the cross-band at 
the apex of the fore wing venation is 
broadly interrupted. The wing pattern be- 
yond the venation in these three species 
is, however, different. In addition, L. ascia 
is distinguished from L. itsta and L. iiirba 
by its larger size, entirely blackish thoracic 
dorsum and abdomen, dark setae on fron- 
tovertex, densely pubescent eyes, and dif- 
ference in the shape of the sculptural cells 
of the scutellum. 

7. Leptotnastidea lamto Prinsloo, new 
species 

(Figs. 4, 26-27) 

Description. — Female. Length: 0.7-1.1 
mm. Colour: Head with frontovertex dark 
yellow from occipital margin to just above 
upper limits of scrobes, lower part of head 
white; setae on frontovertex dark. Anten- 
na, save white apex of pedicel, either en- 
tirely dark brown or with scape white be- 
low along its entire length. Thoracic dor- 
sum variable: more or less concolorous 
with frontovertex or with entire mesono- 
tum, or axillae and scutellum only, with 
brown to blackish-brown suffusions; te- 
gula whitish basally, apical half dark; pro- 
podeum largely blackish-brown; side of 
thorax with prepectus white, mesopleuron 
whitish, with darker suffusions in most 
specimens; mesonotal setae whitish. Fore- 
wing with an oblique dark cross-bands be- 



low submarginal vein; disc from level of 
apex of ventation fuscous with a large 
oblique hyaline area extending from an- 
terior wing margin to near posterior mar- 
gin, as shown in Fig. 4; in sonie specimens 
extreme apex of wing also hyaline; hind 
wing palely infuscated near base and 
along anterior margin beyond venation. 
Legs whitish except middle and hind cox- 
ae blackish-brown, dorsal niargins of all 
femora, and hind tarsus, with brownish 
suffusions in some specimens. Gaster 
whitish with syntergum and apex black- 
ish-brown. Head: 2.0-2.3 X as wide as fron- 
tovertex at median ocellus; ocelli in an ap- 
proximately right-angled triangle, lateral 
pair separated from inner eye margins by 
slightly more than own dianieter; head 
with dimensions and sculpture otherwise 
much as illustrated for L. usta; eyes fairly 
densely but very finely setose, appearing 
naked under low magnification, setae 
about as long the diameter of eye facet; 
frontovertex from median ocellus to occip- 
ital margin with numerous long, dark se- 
tae extending along entire length of each 
inner eye margin in a single row; front as- 
pect of head otherwise finely and sparsely 
setose. Antenna (Fig. 27) with scape about 
5.7 X as long as wide; pedicel ranging 
from slightly shorter to as long as basal 
funicle segment; funicle segments sube- 
qual in length, basal segment about 4X as 
long as broad; club as long as distal two 
funicle segments combined; linear sensil- 
lae present on club and funicle segments 
III-VI. Thorax: dimensions, sculpture and 
setation of mesonotum much as described 
and illustrated for A. usta. Forewing (Fig. 
4) 2.7-3.3 X as long as wide; costal cell 
very narrow, hardly discernible in some 
specimens; venation (Fig. 25) with post- 
marginal vein L5-2.0X kmg as stigmal, 
about 3.3 X the length of the marginal 
vein; disc with setae confined to the large 
hyaline area beyond venation much finer, 
shorter and paler than setae on remainder 
of disc. Ovipositor (Fig. 26): one-third 
length of middle tibia; gonostyli 0.5 X 



160 



Journal of Hymenoptera Research 




Figs. 25-30. Lcptoiiiastidca spp., female paratypes. 25-27. L. laiiito. 25, Apex of forewing venation. 26, Ovi- 
positor and middle tibia drawn to the same scale. 27, Antenna. 28-30. L poiuio. 28, Head, frontal view. 29, 
Antenna. 30, Apex of forewing venation. 



length of middle tibial spur, latter slightly 
shorter than adjacent tarsal segment. 

Male. — Unknown. 

Material Exaiiiuicd. — Female holotype, 
32 female paratypes as follows: CAME- 
ROUN: Nkoemvon, viii.1979, D. Jackson 
(1 female); Victoria Bot. Gardens, 
6.xii.l981, Compton (1 female). GABON: 



Foret de la Mondah, 15-25 km N. Libre- 
ville, 25.xi-3.xii. 87, J.S. Noyes (1 female); 
Foret de Sibang, 5km E. Libreville, 30. xi- 
2.xii. 87, J.S. Noyes (1 female). IVORY 
COAST: Lamto, 6.13N 5.02W, xi.l988, J.S. 
Noyes (holotype, 14 females); Sassandra, 
26.ii-l.iii.1984, M. Matthews (1 female); 
Gagnoa, Antonihio, 2-5.iii.1985, M. Mat- 



Volume 10, Number 2, 2001 



161 



thews (2 females). NIGERIA: Ibadan, Oyo 
St., IITA compound, xi.l987, J.S. Noyes (10 
females). TOGO: 10 km NW Kapalime, 
xii.1988, J.S. Noyes (1 female). Holotype 
and paratypes in BMNH; paratypes in 
SANG. 

Rciunrks. — This species, which is evi- 
dently widespread in West Africa, is sep- 
arated from its congeners by the distinct 
maculation of the forewing, in addition to 
a combination of characters which include 
the generally yellow to brown colour of 
the head and thorax, uniformly dark 
brown antennal flagellum, relatively nar- 
row frontovertex and placement of the 
ocelli, slender funicle segments and short 
ovipositor, as described above. 

8. Leptomastidea pwiido Prinsloo, new 
species 

(Figs. 1, 28-30) 

Description. — Female. Length: 0.8-0.9 
mm. Colour: Head with frontovertex from 
occipital margin to near upper limits of 
scrobes yellow, lower part of head white; 
setae on frontovertex silvery-white. An- 
tenna with radicle brown; scape largely 
whitish, suffused with brown dorsally 
along its entire length, ventral margin 
darkened at apex; pedicel brown with 
apex whitish; funicle segments l-lll 
brown, IV-V whitish, VI brown but slight- 
ly paler that basal three segments; club 
brown. Thoracic dorsum with pro- and 
mesonotum yellow, metanotum whitish; 
propodeum whitish with brownish suffu- 
sions posteriorly; tegula white save slight- 
ly darkened apex; thoracic setation sil- 
very-white; side of thorax white. Forewing 
hyaline except for a pale oblic^ue fuscous 
cross-band below submarginal vein and 
dark patch at apex of venation, as in Fig. 
1. Legs white except middle coxa and tar- 
sal tips dark brown. Gaster with basal 
two-thirds or so white, apical third dark 
brown. Head: (Fig. 28) 1.7x as wide as 
frontovertex at median ocellus; ocelli in a 
right-angled triangle, lateral pair separat- 
ed from inner eye margins by about twice 



their own diameter; head otherwise with 
sculpture and setation much as described 
and illustrated for L. usta. Antenna (Fig. 
29) with scape 4.6 X as long as wide; ped- 
icel subequal in length to basal funicle 
segment; funicle segments subequal in 
length, segment I 3x times as long as 
wide; club as long as distal two and a half 
funicle segments combined; club and all 
six funicle segments with linear sensillae. 
Thorax: dimensions, scultpure and setation 
of mesonotum much as in L. usta. Fore- 
wing (Fig. 1) 2.7-2.8 X as long as wide; 
costal cell narrow in its basal one-third, 
not discernible in apical two-thirds; ve- 
nation (Fig. 30) with postmarginal vein 
1.6X length of stigmal, 3.3-3.5 X as long as 
marginal, latter 0.5 x as long as stigmal; 
wing disc fairly evenly and densely setose 
from base to apex, setation uniform 
throughout, not divided into areas of fine 
and coarse setae as in most other species 
of genus. Ovipwsitor: slightly distorted in 
single slide-mounted specimen, appearing 
about 0.5 X length of middle tibia, gonos- 
tyli 0.5 X as long as middle tibial spur, lat- 
ter subequal in length to basal tarsal seg- 
ment of niiddle leg. 

Male. — Colour: Head and thorax much 
as in female; antenna with scape largely 
whitish, otherwise entirely dark brown; 
forewing patterned as in female; legs en- 
tirely whitish except dark tarsal tips, mid- 
dle coxa very slightly embrowned; gaster 
entirely dark brown. Structure: Differing 
from female mainly in toruli which are 
placed higher on face, eyes which are 
slightly smaller and antennal shape: an- 
tenna with funicle segments subequal in 
size, tapering strongly at their apical ends, 
basal segment 5x times as long as broad; 
funicle clothed with whorls of long, 
curved setae, each seta approximately 5x 
as long as the width of a segment; club as 
long as distal two funicle segments com- 
bined with a longitudinal row of four 
spine-like setae near base. Phallobase one- 
third length of middle tibia in single slide- 
mounted paratype. 



162 



Journal of Hymenoptera Research 



Material examined — Female holotype, 5 
female, 3 male paratypes (SANC) as fol- 
lows: SOUTH AFRICA: KwaZulu-Natal 
Province: Port Edward, i.l972, H.P. Insley, 
ex mealybugs on Maytenus itudata 
(Thunb.) Blakelock (T 4166). 

Remarks. — Leptomasiidea pondo, which is 
known only from South Africa, differs in 
the female from other species of the genus 
by the forewing which is mostly hyaline 
except for the presence of a cross-band be- 
low submarginal vein and dark patch at 
apex of venation. It can be distinguished 
further from its Afrotropical congeners in 
the female by the generally pale yellow 
appearance, white hind coxa, broad fron- 
tovertex, and uniformly setose forewings. 

ACKNOWLEDGEMENTS 

I thank j.S. Noyes (BMNH) and J.L. Nieves Aldrey 
(MNCN) for the loan of specimens and my colleague 
Ottilie Neser for technical assistance and reading the 
manuscript. The digital photographs of the wings 
were taken by Alan Hall, University of Pretoria, and 
Elsa van Niekerk, PPRl, kindly assisted in preparing 
them for publication. 

LITERATURE CITED 

Compere, H. 1939. A second report on some miscel- 
laneous African Encyrtidae in the British Muse- 
um. Bulletin of Eiitoiiiological Research 30: 1-26. 



Girault, A. A. 1915. Four new encyrtids from Sicily 
and the Philippines. E)ito}iiologist 48: 184-186. 

Mercet, R. G. 1916. Chalcfdidos de Espana. Boletiii de 
In Real Sociedad Espauola de Historia Natural 16; 
112-117. 

Mercet, R. G. 1924. Los generos Leptoiiiastidea, Callip- 
tenviia y Gi/niiiiisa. Bolctiu de In Renl Sociedad Es- 
pninola de Historia Natural 24: 252-260. 

Noyes, J. S. 1981. On the types of the species of En- 
cyrtidae described by R. Garcia Mercet (Hyme- 
noptera: Chalcidoidea). Eos, Madrid 55/56: 165- 
189. 

Noyes, J. S. 1988. Encyrtidae (Insecta: Hymenoptera). 
Fnuiin of New Zealand 13: 1-188. 

Noyes, J. S. 2000. Encyrtidae of Costa Rica (Hyme- 
noptera: Chalcidoidea), I. Memoirs of the American 
Entomological Institute 62: 1-355. 

Noyes, J. S. and M. Hayat. 1994. Oriental meahfbug 
parnsitoids of the Anagyri)ii {Hymenoptera: E)jci/rti- 
dae). CAB International, Wallingford, UK, i-viii + 
554 pp. 

Prinsloo, G. L. 1983. The southern African species of 
Gifranusoidea Compere (Hymenoptera: Encyrti- 
dae). journal of the Ento))iological Society of South- 
ern Africa 46: 103-113. 

Timberlake, P. H. 1918. New genera and species of 
Encyrtidae from California parasitic in mealy- 
bugs (Hymenoptera). U)iiversiti/ofCnlifornin Pub- 
licatio)is in Entomology 1: 347-367. 

Trjapitzin, V. A. 1989. Parasitic Hymenoptera ofthefnm. 
Encyrtidae of Palaearctics. Opredeliteli po faune 
SSSR, Izdavayemyie Zoologicheskim institutom 
AN SSSR 158. Leningrad, Nauka, Leningrad Di- 
vision, 488 pp. (in Russian). 



]. HYM. RES. 
Vol. 10(2), 2001, pp. 163-180 

Investigation of the Biology of Hymenoptera Associated with 

Fergitsonina sp. (Diptera: Fergusoninidae), a Gall Fly of 
Melaleuca qiiinqiienervia, Integrating Molecular Techniques 

J. A. GOOLSBY, C. J. BURWELL, J. MaKINSON, AND F. DRIVER 

(JAG) United States Department of Agriculture, Agricultural Research Service, Australian 

Biological Control Laboratory, CSIRO Long Pocket Laboratories, 120 Meiers Rd. Indooroopilly, 

QLD, Australia 4068 (email: iohn.goolsby@brs.ento.csiro.au); (CJB) Queensland Museum, P.O. 

Box 3300, South Brisbane, QLD, Australia 4101; (JM) Commonwealth Scientific and hidustrial 

Research Organisation (CSIRO), Division of Entomology, Australian Biological Control 

Laboratory, 120 Meiers Rd. Indooroopilly, QLD, Australia 4068; (FD) CSIRO Entomology, Black 

Mountain Laboratories, GPO Box 1700, Canberra, ACF, Australia 2601 



Abstract. — The biologies of eleven species of Hymenoptera associated with the multi-locular 
galls of the fly, Fergtison!)ui sp. (Fergusoninidae) were investigated. More than 2000 wasps were 
reared from 1100 galls collected in Queensland and New South Wales, Australia over a two-vear 
period from 1997 to 1999 from Melaleuca qiiiuqiiciicrvia (Myrtaceae). Additional galls from each 
site were dissected for observation and description of the immature stages. A molecular technique, 
which involved sequencing the D2 expansion domain of the 28S rRNA gene, was used to match 
the identity of the immature wasps with their adult forms. Of the eleven species of Hymenoptera 
associated with the Fergiisonina sp. galls, we were able to observe and describe the biology of 
nine of the species. Eiin/tonia sp., Cocloci/ba sp, Neanastatus sp., Cirwspiliis sp., Bracoii sp., Mcgas- 
tigDius sp. and Poecilocri/pitiis uignviiaciilatiis Cameron, commonly or exclusively, fed directly upon 
the Fergiisonina larvae and or pupae with most species developing on a single host. However, 
Eurytoma sp., Bracon sp., and P. nigroiiiaciilatiis usually fed on multiple hosts. These species have 
specialized biologies, which enable them to chew through plant tissues to access gall inhabitants. 
Chromeim/tivna sp. and Eiuieriis sp. appeared to be hyperparasitoids based on the available e\i- 
dence. The biological control implications of this suite of Hymenoptera are discussed in terms of 
their regulatory effect on Fergiisonina sp., a potential biological control agent of M. qiiinqiienervia, 
an invasive weed in Florida, USA. 



Species of Fergusouinn (Fergusoninidae) (QLD). The gall-making cyclorrhaphous 

and their associated Fcrgusobiii nematodes fly is under study as a pc^tential biological 

(Tylenchida: Sphaerulariidae), together control agent for M. quiiiquciicrvia, which 

form galls on the buds of their myrtaceous was introduced from Australia into Flori- 

host plants (Currie 1937, Ferrar 1987, Gi- da in the United States in the early 1900's. 

blin-Davis 2000). An undescribed species In the last 30-40 years M. qiiiiiqiioicrvin 

of Fergiisonina and an undescribed Fcrgii- has greatly expanded its range in southern 

sobin form vegetative and floral galls on Florida, including the ecologically sensi- 

the broad-leafed paperbark tree, Melaleuca Ave Everglades, where it now infests over 

qiiinqiienervia (Cavanilles) S.T. Blake (Fig 200,000 hectares causing extensive envi- 

1). This Fergiisonina sp. is present through- ronmental and economic damage (Turner 

out the Australian distribution of M. (/;////- et al. 1998). 

quenervia, which stretches along the east The seasonal phenology of Fergiisonina 

coast from southern New South Wales sp. on M. quinqiienervia was investigated 

(NSW), to the far north of Queensland by Goolsby et al. (2000a) over a two-year 



164 



Journal of H^ menoptera Research 




Fig 1. Fcrgiisoiiiiin sp. gall on Mclnlfiicn qiiiiiqiioicrvia; a, Intact gall showing cluster of chambers, some with 
exit holes, and b, cross-section of gall showing individual chambers of the flv larvae. 



period between 1997 and 1999. The study 
indicated that biotic factors, including par- 
asitism, might have a significant effect on 
Fergusoiiina sp. gall density. Data collected 
for emergence of flies and associated gall 
inhabitants revealed numerous Hymenop- 
tera, comprising nine species of Chalci- 
doidea and a single species each of Bra- 
conidae and Ichneumonidae, all potential 
parasitoids of the Fcrgnsonina sp. In order 
to understand the impact of parasitism, 
we first needed to establish which species 
of wasps are primary parasitoids of the 
Fergusoiiiiin sp., which are hyperparasi- 
toids or inquilines and which can behave 
as both primary parasitoids and hyper- 
parasitoids. A large complement of pri- 
mary parasitoid species may indicate that 
parasitism plays a significant role in reg- 
ulating Fcrgusojiiiin sp. populations in 
Australia. Understanding the regulatory 
effect of natural enemies on a potential bi- 
ological control agent in its native range is 
a useful predictor of its success in its ad- 
ventitious range. 

In his pioneering work on the Ferguson- 
inidae, Currie (1937) postulated that par- 
asitic Hymenoptera played a major role in 
regulating their population dynamics. He 
reared niany species of wasps from flower 



bud galls produced by Fcrgiisoiiiim iiichol- 
soiii Tonnoir on Eiicah/pHiis iiincrorhi/iicha F. 
Mueller ex. Bentham and dissected galls to 
determine the biologies of their imma- 
tures. Currie noted that both a chalcidoid 
and a braconid were true parasitoids of 
the gall-making flies and briefly listed four 
other species of chalcidoid wasps that 
formed independent chambers within the 
galls. However, more detailed information 
on the biologies of the gall-associated 
wasps was never published. Taylor ct al. 
(1996) reared twelve species of wasps 
from leaf galls formed by Fcrgusoiiiiui flav- 
iconiis Malloch on Eitcnh/ptits cmimlduleusis 
Dehnhardt in South Australia. They did 
not dissect galls, but discussed the prob- 
able biologies of the various wasp species 
in the light of their relative abundance and 
the biologies of related species. Both stud- 
ies found an abundance of gall-associated 
Hymenoptera, but were largely unable to 
positively determine their role inside the 
gall. 

Gall-making agents interact with asso- 
ciated parasitoids, predators and inqui- 
lines behind the cover of plant tissue that 
often obscures our understanding of their 
biologies. Because it may be difficult to 
identify the hymenopteran larvae associ- 



Volume 10, Number 2, 2001 



165 



ated with galls, many studies fail to asso- 
ciate the biology of the immatures with 
their adult form (Shorthouse et al. 1990, 
Manongi and Hoffman 1995). The most 
common method for determining the bi- 
ology of immatures is to observe theni in 
the gall and then hold them until they 
emerge as adults, which can be more eas- 
ily identified. This method is the most 
straightforward and has been used widely 
in the study of gall inhabiting Hymenop- 
tera. 

However, this method is time consum- 
ing and may not be practical when dealing 
with galls that contain a large suite of par- 
asitoid species. In our study we also dis- 
sected and observed gall contents, but 
combined this method with a molecular 
technique which involved sequencing the 
D2 expansion domain of the 28S rRNA 
gene to match the identity of the wasp lar- 
vae with their adult forms. The D2 expan- 
sion domain of the 28S rRNA gene has 
been used in other studies to separate 
cryptic species of adult hymenopteran 
parasitoids (De Barro et al. 2000, Babcock 
and Heraty 2000), and aquatic weevils 
(Goolsby et al. 2000b). Tilmon et al. (2000) 
used the COl gene to determine species 
composition of immature parasitoids in 
their host. We used the molecular method 
of sequencing the D2 gene as a way to de- 
termine identity of the immatures as we 
observed their biology /;/ vivo. 

MATERIALS AND METHODS 

Monthly collections of mature Fergusoi- 
iiia sp. galls on A/I. quifiqiieiiervia trees were 
made from Peregian and Morayfield 
(QLD) and Woodburn (NSW) from July 
1997 to September 1999. The locations and 
phenology of the Fergusouiria sp. are de- 
scribed in Goolsby et al. (2000a). Galls 
were held for one month in ventilated 
containers for emergence of the gall inhab- 
itants. The emerged insects were counted 
and sorted to species. 

In September 1999, following the two- 
year study, approximately 30 galls were 



collected from each site in order to ob- 
serve and investigate the biology of the 
gall inhabitants. We dissected several hun- 
dred gall chambers in order to observe the 
behavior of the gall inhabitants. Observa- 
tions of the gall insects were made using 
a dissecting microscope, and pictures of 
the contents were taken using a digital 
camera (Sony Mavica, model FD-88). Ow- 
ing to the mobility of the camera, pictures 
of immatures could be taken by focusing 
through the ocular of the microscope. Im- 
matures were placed in vials of 95% al- 
cohol for DNA analysis. Several speci- 
mens of each species were analyzed. 
Adult parasitoids were identified to genus 
and, where possible, to species. Vouchers 
are located in the Queensland Museum, 
Brisbane; Florida State Collection, Gaines- 
ville and the U.S. National Museum, 
Washington, D.C. 

Eggs, larvae, and pupae of Hymenop- 
tera were used for gene sequencing. Gene 
sequences of the immature Hymenoptera 
were compared with adults that had been 
reared from Fergusouiua sp. galls. Adult 
representatives of the less common Hy- 
menoptera species were reared from Fer- 
gusouiua sp. galls collected during the pre- 
vious two years. We sequenced the D2 ex- 
pansion domain of the 28S rRNA, which 
ranged from 564 to 593 base pairs long de- 
pending on the species. The methods were 
those described by De Barro et al. (2000). 

The polymerase chain reaction (PCR) 
was used to amplify the D2 gene regions 
for each specimen. Primers for the region 
followed Campbell et al. (1993); D2F 5'- 
CGTGTTGCTTGATAGTGCAGC-3' and 
D2R 5'-TTGGTCCGTGTTTCAAGACGG-3', 
or ND2F 5'- AGTACCGTGAGGGAAA- 
GTTG-3', which was used in some reac- 
tions as an alternate forward primer 
which anneals approximately 90 bases 
down-stream of the D2F binding site. All 
reaction volumes were 50 fxl, containing 
20 pM of each primer, 200 txM each dGTP, 
dATP, dCTP and dTTP, 1.5-2.5 mM 
MgCU, 2 |jl1 DNA lysate, IX supplied buff- 



166 



JOURNAL OF HYMENOPTERA RESEARCH 



Tnhle 1. Gall insects reared during two-year field study. 



Mor.ntiL 
4^3 



Woodburn 
263 



All Mtf!, 

1122 



Fci'i^iisoiiiiHi sp. 


372 


483 


420 


1275 


Enn/tonm sp. 


300 


473 


13 


786 


Coclocyha sp. 


276 


33 


122 


431 


Ncaiiastatiis sp. 


65 


144 


35 


244 


Cirwspilus sp. 


113 


19 





132 


Bnicoii sp. 


28 


42 


47 


117 


Eiipcliinis sciuipiitata 


4 


99 





103 


Chwuicuriftoiiin sp. 


15 


50 


5 


93 


Mt'gastii^fiins sp. 


17 


35 


28 


80 


Eiipehmis (Eiipchinifi) sp. 


3 


5 


2 


10 


Eudcrus sp. 


3 


4 


2 


9 


Poccilocn/ptus lugnviinciilatiis 





1 


4 


5 


Total Hymenoptera 


824 


905 


258 


2010 


Jo Parasitism 


68.90% 


65.20'X, 


38.05% 


61.19% 



er and 2.5 U Taq polymerase (Bresatec, 
Australia). PCR amplification was done 
using a Hybaid thermocycler using the 
following parameters. A pre-cycle dena- 
turation step for 5 min at 94°C, followed 
by the addition of the Taq polymerase. 
Then, 35 cycles of 1 min at 94°C, 1 min at 
55°C and 1.5 min at 72°C followed by a 
final post-cycle extension step at 72°C. 

The D2 amplicons were purified and 
prepared for sequencing by electrophore- 
sis in 0.8% TAE agarose gels containing 10 
fxg ml ' ethidium bromide (Sambrook ct 
al. 1989). Fragments were excised and 
transferred to a microfuge tube. The aga- 
rose slices were mashed in 30 |xl sterile 
distilled water using a toothpick, then in- 
cubated at 50°C for 1 h. Samples were left 
at rc^om temperature overnight to allow 
the DNA to elute from the gel. The sam- 
ples were stored at -20°C until required. 

Five microliters of the eluted PCR-am- 
plicons and the appropriate PCR-primers 
were used for sequencing according to the 
ABI PRISM Dye Terminator Cycle Se- 
quencing Ready Reaction Kit Manual (PE- 
Applied BioSystems). Both strands of each 
fragment were sequenced and reactions 
were loaded onto an Perkin Elmer, Ap- 
plied Biosystems Division, Model 377 ABI 
PRISM Genetic Analyzer. All sequences 
were deposited in GenBank"" (see Table 2). 



RESULTS AND DISCUSSION 

Eleven species of Hymenoptera were 
reared from the Fcrgusonina sp. galls over 
the two-year sampling period (Table 1). 
Euryioina sp. was the most common spe- 
cies at Peregian and Morayfield, whereas 
Coclocyha sp. was most common at Wood- 
burn. Eiin/tonia sp. and Coclocyha sp. com- 
prised 61% of the 2010 specimens of Hy- 
menoptera reared froni the galls over the 
two-year period. Parasitism of the ¥cr- 
giisoiiiiia sp. larvae and pupae was high 
(> 60%) at both Morayfield and Peregian. 
The pooled percentage parasitism for the 
three sites over the two year period was 
61.2% (Table 1). 

We identified the immatures of eight of 
the eleven species collected in the study 
by matching their DNA sequences with 
those of the adult forms (Table 2). The lar- 
vae of a ninth species were identified by 
examining larval exuviae recovered from 
Fcr^usoiiiiia puparia from which parasit- 
oids had emerged. Immatures of the re- 
maining two species were not encoun- 
tered; their biology was deduced from 
published informaticin of congeners. All of 
the immatures analyzed were matched 
with adult forms except for one hyperpar- 
asitoid egg. The D2 sequence from the hy- 
perparasitoid egg was unique and not de- 



Volume 10, Number 2, 2001 



167 



Table 2. Gall insects identified in study using D2 sequence data. 



Species* 



Stage 



Selection criteria 



1 


Fcr\;iisoi!iiin-Ma\e 


MORAYFIELD 


Adult 


AF345569 


2 


Fergusoiii}ia-¥emale 


WOODBURN 


Adult 


AF345570 


3 


Fcrgusoiiiita 


MORAYFIELD 


Larva 


AF345571 


4 


Eiinftoiiia 


MORAYFIELD 


Adult 


AF345572 


5 


Eiin/toiim 


PEREGIAN 


Adult 


AF345573 


6 


Eun/toiiia 


PEREGIAN 


Adult 


AF345612 


7 


Eiiryfpnia 


MORAYFIELD 


Larva 


AF345574 


8 


Eiiri/toiua 


PEREGIAN 


Larva 


AF345575 


9 


Eiirytoiiia 


MORAYFIELD 


Larva 


AF345613 


10 


Eiin/toiim 


MORAYFIELD 


Larva 


AF345606 


11 


Eiin/toiua 


PEREGIAN 


Larva 


AF345610 


12 


Eiiri/toiiia 


MORAYFIELD 


Larva 


AF345607 


13 


Hyperparasitoid egg 


MORAYFIELD 


Egg 


AF345608 


14 


Cocloci/bn 


PEREGIAN 


Adult 


AF345576 


15 


Ciuioci/ba 


PEREGIAN 


Adult 


AF345577 


16 


Cocloci/ba 


PEREGIAN 


Adult 


AF345614 


17 


Coi'locyba 


PEREGIAN 


Adult 


AF359455 


18 


Coi'locifba 


WOODBURN 


Larva 


AF359454 


19 


Ncanastntus 


MORAYFIELD 


Adult 


AF345580 


20 


Neanastntiis 


PEREGIAN 


Adult 


AF345582 


21 


Ncniiastatiis 


MORAYFIELD 


Pupa 


AF345581 


22 


Ncanastntus 


PEREGIAN 


Larva 


AF345583 


23 


Ncaiiastatiis 


MORAYFIELD 


Larva 


AF345584 


24 


Ncanastntus 


MORAYFIELD 


Larva 


AF345615 


25 


Cinospilus 


PEREGIAN 


Adult 


AF345585 


26 


Cinvspilus 


MORAYFIELD 


Adult 


AF345586 


27 


Cirwspilus 


MORAYFIELD 


Adult 


AF345616 


28 


Cin-ospilus 


PEREGIAN 


Larva 


AF345604 


29 


Cinvspilus 


PEREGIAN 


Larva 


AF345605 


30 


Brncon 


WOODBURN 


Adult 


AF345587 



Primary gall former 

Primary gall former 

Primary gall former 

Reared from individual that had predat- 
ed on multiple hosts 

Reared from individual that had predat- 
ed on multiple hosts 

Reared from pupa in chamber with copi- 
ous masticated gall tissue and black 
meconium 

Larva had fed on multiple hosts, cham- 
ber contained ball of Fcr;^uso)una re- 
mains 

Larva had fed on multiple hosts, copious 
amounts of masticated gall tissue pre- 
sent 

Larva collected at center of young gall 
with connections to two other cham- 
bers 

Larval remains being consumed by Brn- 
con larva 

Larva with Poccilocn/ptus larva attached 
(Specimen 53) 

Fcrgusiviina parasitoid in clean gall 
chamber, attacked by hyper-parasitoid 
below (Specimen 13) 

Egg attached to Eun/tonm larva (Speci- 
men 12 above) 

Reared from naked pupa, chamber con- 
tained Fcrgusonina remains 

Reared from an intact Fcrgusoiuun pupar- 
ium 

Reared from an intact Fcrgusoninn pupar- 
ium 

Reared from an intact Fcrgusoninn pupar- 
ium 

Larva restricted to single chamber 

Reared from Fcrgusoninn gall 

Reared from Fcrgusoninn gall 

Naked pupa restricted to single chamber 

Larva collected in single chamber, cra- 
dling remains of Fcrguso)rina 

Same as above, but from different site 

Larva collected from single isolated 
chamber 

Reared from Fcrgusoniiui gall 

Reared from Fcrgusoninn gall 

Reared from Fcrgusomnn gall 

Collected from intact chamber with rem- 
nants of Fcrgusoninn 

Collected frorr^ intact chamber with rem- 
nants of Fcrgnsonnm 

Reared from silk cocoon inside Ferguson- 
iim gall 



168 



Journal of Hymenoptera Research 



Table 2. Continued. 





Species* 




Locafion 


Stage 


Con Bank" 
accession 
number 


Selection criteria 


31 


Bmcoii 




MORAYFIELD 


Adult 


AF345588 


Reared from Fergusoiiiua gall 


32 


Bmcoii 




MORAYFIELD 


Larva 


AF345589 


Larva collected as it was making silk co- 


33 


Bracon 




MORAYFIELD 


Larva 


AF345590 


coon 

Larva collected as it straddled two 
chambers, both full of copious masticat- 
ed gall tissue and Fergusouirm remains. 


34 


Bracoii 




MORAYFIELD 


Larva 


AF345617 


Larva collected from gall with intercon- 
nected chambers, full of masticated gall 


35 


Eiipcbiius sciiiipiitnta 


MORAYFIELD 


Adult 


AF345591 


tissue and Fergusouirm remains 
Reared from Fergusoniria gall 


36 


Eupclnius ficiitipi 


utata 


MORAYFIELD 


Adult 


AF345592 


Reared from Fergusorriun gall 


37 


Eupt'hmis i^ciniputnta 


MORAYFIELD 


Adult 


AF345618 


Reared from Fergusorriun gall 


38 


Cliwnwuri/toiim 




PEREGIAN 


Adult 


AF345595 


Found in single isolated gall chamber 


39 


Chnvnciuytoina 




MORAYFIELD 


Adult 


AF345596 


Reared from Fergusorriun gall 


40 


Chroineurytoma 




MORAYFIELD 


Adult 


AF345620 


Reared from Fergusorriun gall 


41 


Chroiueun/toma 




MORAYFIELD 


Larva 


AF345597 


Collected front chamber with Nenirastntus 


42 


Chmiiiciin/toiim 




MORAYFIELD 


Larva 


AF345598 


pupal remains 

Collected from chamber with Nearrnstntus 

pupal remains 


43 


Chroinciiri/tODia 




MORAYFIELD 


Larva 


AF345593 


Larva collected from intact chamber with 


44 


Mi\^astignins 




MORAYFIELD 


Adult 


AF345594 


wasp pupal remains 

Reared from Fergusorriun pupal case 


45 


Megastig)nns 




WOODBURN 


Adult 


AF345619 


Reared from Fergusorriun gall 


46 


Eupehnus (Eupc, 


Innis) 


MORAYFIELD 


Adult 


AF345599 


Reared from Fergusorriun gall 


47 


EupeUniis (Eupci 


btuis) 


PEREGIAN 


Adult 


AF345600 


Reared from Fergusorriun gall 


48 


Eupehnus {Eupci 


Iduis) 


PEREGIAN 


Adult 


AF345621 


Reared from Fergusouirm gall 


49 


Euderus 




PEREGIAN 


Adult 


AF345601 


Reared from Fergusonirra gall 


50 


Euderus 




WOODBURN 


Adult 


AF345602 


Reared from Fergusouirm gall 


51 


Euderus 




MORAYFIELD 


Egg 


AF345609 


Egg collected from deflated, dead Poecilo- 
cryptus larva 


52 


Poecilocryptus u, 
culatus 


igroiiia- 


WOODBURN 


Adult 


AF345603 


Adult collected from Fergusorriim gall 


53 


Poecilocryptus u, 
culntus 


igwiiui- 


PEREGIAN 


Larva 


AF345611 


Hyperparasitoid of Eurytorrm larva (Spec- 
imen 11) 



No variation was noted in D2 gene sequence data between individuals of each taxon. 



tected again. Most species develop in a 
single intact gall chamber as primary par- 
asitoids of Fergusoiiiiia sp. The larvae of 
three species, Eiiiytouin sp., Bracoti sp. and 
Poecilocryptus nigromaculntus tunnelled be- 
tween gall chambers to feed on multiple 
hosts. Some authors (eg. Godfray 1994) 
prefer the term predator to describe this 
biology. We consider the distinction be- 
tween predator and parasitoid to be am- 
biguous and prefer to describe these spe- 
cies as parasitoids feeding on multiple 
hosts. Several hyperparasitoid species 



were identified either by their eggs, that 
were found attached to parasitoid larvae, 
or by their larvae that occupied chambers 
containing the remains of parasitoid lar- 
vae or pupae. 

Biology of Associated Hymenoptera 

Eurytonia sp. (Eurytomidae). — Speci- 
mens of Eurytoma emerged from galls 
from all three sites and made up 39.170 of 
the total number of Hymenoptera reared 
from the field collections. They were the 
most numerous gall-associated wasps at 



Volume 10, Number 1, 2001 



169 



the two Queensland localities, Peregian 
and Morayfield. Few Eurytouia were re- 
covered from galls collected at Woodburn 
(Table 1). Despite some variation in the 
coloration of their legs, all specimens ap- 
pear to belong to a single species. The D2 
gene sequences of adults from Peregian 
and Morayfield were identical (Table 2). 

Eiiryionia is an enornious cosmopolitan 
genus; Boucek (1988) listed 66 species 
from Australia and indicated there were 
many still undescribed. In the absence of 
revisionary studies on Australian species 
of Euryiouia it was not possible to identify 
this species. However, it appears to be al- 
lied to a distinctive group of Australian 
Eurytouin discussed by Boucek (1988) and 
characterized mainly by an elongate peti- 
ole and relatively long marginal vein. This 
group includes £. lougipetiolntn Girault and 
£. austrnlieusis Ashmead (Boucek 1988) 
and after examination of their types by 
CJB, we believe that £. carlylci Girault and 
E. herberteiisis Girault probably also be- 
long here. The species of Eurytonui reared 
from Fergusouina galls possesses several 
characters of the group including an elon- 
gate female petiole that is longer than the 
hind coxae and slightly curved, and a lat- 
erally compressed gaster that has the com- 
bined length of the first three gastral ter- 
gites less than their height and shorter 
than the length of the fourth gastral tergite 
(Boucek 1988). However, species of the 
group have the marginal vein 2.5-3 X lon- 
ger than the stigmal vein (Boucek 1988) 
while the Eurytoma sp. from the galls has 
the marginal vein only about twice the 
length of the stignial. 

Eurytoma sp. larvae had a relatively 
large head capsule with the mouthparts 
directed ventrally. The mandibles of ma- 
ture larvae were bidentate with a strong, 
acute, apical tooth and an acute subapical 
tooth about half the length of the apical. 
Larvae had a series of conspicuous dorsal 
protuberances on the meso- and metatho- 
rax and the first eight abdominal seg- 
ments. Larvae were only moderately se- 



tose with most thoracic and abdominal se- 
tae short and inconspicuous. However, 
each thoracic segment had three pairs of 
longer setae ventrad of the spiracles. 

The larval biology of Eurytoma sp. was 
variable. Commonly lan.'ae fed on multi- 
ple Fergusouina larvae and possibly the 
larvae of other gall-associated Hymenop- 
tera. Occasionally individual Eurytoma pu- 
pae were observed in single intact cham- 
bers, having completed their development 
on only one Fergusouina host. Molecular 
data confirmed that Eurytoma developing 
on single or multiple hosts were the same 
species. 

Eurytoma sp. larvae feeding on multiple 
hosts were found in chambers that were 
connected by small holes to one or more 
other chambers. These chambers were 
typically filled with brown, particulate de- 
bris that we interpreted as masticated, but 
not ingested, gall tissue. In addition, dis- 
sociated plates from the dorsal shields of 
Fergusouina larvae were found amongst 
the brown debris. Frequently several Eur- 
ytoma larvae completed development in a 
single gall. 

Eurytoma is a diverse genus with a wide 
array of larval biologies ranging from en- 
tomophagous to phytophagous species. 
Many species attack gall formers (Di Gui- 
lio 1997) including some that feed on sev- 
eral hosts in multi-chambered galls (Blair 
1944, Boucek 1988), which is similar to the 
species in our study. The larvae of some 
gall-inhabiting species of Eurytoma have 
been reported to feed on both insect and 
plant tissue (Varley 1937, Noble 1941, 
Askew 1961). Although the Eurytoma lar- 
vae in our study masticated gall tissue, it 
is not clear if they derived any nutritional 
benefit from this activity, or if they just 
mechanically scraped away the tissue to 
gain access to additional chambers. 

All studies on the Hymenoptera asso- 
ciated with Fergusouina galls have record- 
ed Eurytoma species (Currie 1937, Harris 
1982, Taylor et nl 1996) but only Currie 
investigated the lan'al biology. Unlike our 



170 



Journal of Hymenoptera Research 



study, he found that larvae of Euri/toma 
"varirufipes" (an unpublished Girault 
manuscript name), were inquilines within 
galls of Fergiisoiiiiin nicholsoui Tonnoir on 
Eucalyptus macrorhyndia, forming separate 
chambers to those of the fly larvae. 

Coelocyba sp. (Pteromalidae). — Overall, 
Coelocybn sp. was the second most abun- 
dant wasp reared (21.4% of total speci- 
mens), but was uncommon at the Moray- 
field site (Table 1). Boucek (1988) listed 
nine species of the endemic Australian ge- 
nus Coelocyba, which he noted was com- 
posed of two species groups separated on 
the structure of the dorsellum and pro- 
podeum. The species reared from Fergu- 
soniiia sp. galls belongs tc^ the group con- 
taining C. nigrocincta Ashmead, character- 
ized by the posterior margin of the dor- 
sellum being broadly rounded (Boucek 
1988). Coelocyba sp. closely resembles C. 
uigrocinctn in color pattern, however spe- 
cies in the genus are difficult to recognize 
and the value of color in distinguishing 
species is questionable (Boucek 1988). The 
genus is in need of revision (Boucek 1988) 
and consequently precise identification of 
the species reared in the study was not 
possible. 

Coelocyba larvae appear almost gla- 
brous, distinguishing them from most lar- 
vae encountered during dissections, ex- 
cept those of Cirrospilus. Coelocyba larvae 
can be distinguished from Cirrospilus lar- 
vae by their globular head capsules and 
ventrally directed mouthparts. In addi- 
tion, Coelocyba larvae have tridentate man- 
dibles with a strong, acute apical tooth; a 
closely appressed, acute, subapical tooth; 
and a small basal tooth. 

Adults of Coelocyba sp. emerged either 
from naked wasp pupae or from intact 
Fergusouiua puparia in approximately 
equal proportions. The D2 gene sequences 
of adults reared from both were identical 
(Table 2). Coelocyba that emerged from Fer- 
gusonina puparia either developed as true 
endoparasitoids or more probably as ec- 
toparasitoids of the pharate Fergusoiiiiia 



pupa. Chambers that contained parasit- 
ized Fergusonina puparia closely resem- 
bled those with unparasitized puparia. 
Parasitized puparia were attached to the 
wall of the chamber by the normal trans- 
parent, elastic substance (see Currie 1937: 
150). Naked pupae and larvae of Coelocyba 
were always found singly in isolated 
chambers, along with pale granules of 
host remains containing Fergusonina dor- 
sal plates. In these cases Coelocyba devel- 
oped as a primary ectoparasitoid of the 
Fergusonina. We found no evidence that 
Coelocyba larvae fed on gall tissue. Cham- 
bers with Coelocyba pupae also contained 
a patch of dark meconium. 

Known species of Coelocyba are associ- 
ated with gall-inducing pteromalids and 
fergusoninids (Boucek 1988), but their pre- 
cise larval biologies are unknown. Boucek 
(1988) and Taylor et al. (1996) reported 
that larvae of C. nigrocincia Ashmead had 
been demonstrated to be inquilines in the 
galls of Perilampella hecataeus (Walker) on 
Acacia decurrens Willdenow, primarily 
based on work done by Noble (1941). 
They claimed that the C. nigrociiicta larva 
killed the resident gall-inducer and 
formed its own cell and fed on the gall 
tissue. However, although Noble (1941) 
reported that the C. nigrociiicta larva killed 
the larva of the gall-inducer, he made no 
mention of it forming its own chambers, 
or of it feeding on gall tissue. In our study 
we found no evidence that Coelocyba sp. 
larvae fed on gall tissue and concluded 
that they were almost always primary par- 
asitoids of the Fergusonina larvae or pu- 
pae. This is in agreement with Currie 
(1937) who briefly noted that the species 
of Coelocyba that he reared from flower 
bud galls produced by Fergusonina nichol- 
soni on Eucalyptus macrorhyncha was a 
"true parasite" of the fly larvae. Taylor et 
al. (1996) reared C. nigrocincta from leaf 
bud galls on £. macrorhyncha but did not 
investigate its larval biology. 

Neanastatus sp. (Eupelmidae). — A single 
species of Neanastatus was moderately 



Volume 10, Number 2, 2001 



171 



common at all three sites (Table 1) and ac- 
counted for 12.1% of the total specimens 
reared. Molecular sequences of adults 
from Morayfield and Peregian were iden- 
tical (Table 2). Species of Ncaiiastatiis are 
apparently restricted to the Old World 
(Gibson 1989), but are widely distributed 
fron"i southern Europe through Africa, and 
southern Asia to Australia with most spe- 
cies known from the tropics (Boucek 
1988). Boucek (1988) lists 21 Australian 
species, all ciescribed by A. A. Girault. 
Specimens reared from galls of Fergnsoni- 
im sp. on M. qiiiuqucucrvia have the head 
and most of the meso- and metasoma 
dark-colored, mostly with metallic green 
reflections. At least the anterior half of the 
pronotum and most of the first gastral ter- 
gite is yellowish. The hind tibia is mostly 
black with a narrow, basal, white band. 
Amongst the Australian species, they 
most closely resemble N. flaviprouotuui Gi- 
rault. However the holotype of this spe- 
cies differs in that the lower face sur- 
rounding the mouthparts is yellowish. In 
addition, the pronotum is extensively yel- 
lowish with only a narrow posterior dark 
band. 

Ncniiiistntiis sp. larvae are whitish and 
fusiform, gradually tapering posteriorly. 
The larval mandibles are simple, each 
with a single acute tooth. Neniinstntiis lar- 
vae are conspicuously setose, with rows of 
long lateral setae on the thoracic and ab- 
dominal segments, except the first abdom- 
inal segment. The thoracic segments have 
two additional pairs of long setae. The lar- 
vae can be distinguished from the setose 
larvae of Oiroineiiiytoiun (see below) by 
the absence of ventrolateral setae on the 
abdominal segments. Neniinstntiis sp. pu- 
pae can be distinguished by a conspicuous 
tubercle on the dorsal frt^ns. 

The available evidence indicates that 
Neannstntus sp. develops as a solitary, pri- 
mary ectoparasitoid of Ft'rgiisoiiiiin larvae. 
In all instances, Neannstntus larvae, pupae 
and newly eclosed adults were found sin- 
gly inside isolated, intact chambers. Ma- 



ture larvae were observed resting on their 
dorsal surfaces and cradling, on their ven- 
tral surfaces, small balls of tissue contain- 
ing Fergiisoiiinn dorsal plates. There was 
no indication of Neanastatus larvae feeding 
on gall tissue. Neanastatus pupae occupy 
relatively clean chambers that contain a 
dark patch of tar-like larval meconium; 
one or sometimes two shriveled, setose, 
larval exuvia; and usually the remains of 
a Fergusoiiina larva indicateci by the pres- 
ence of fragments of the dorsal shield. 

Species c^f Neanastatus have been record- 
ed as parasitoids in the galls of cecido- 
myiid flies, especially those associated 
with grasses and herbaceous plants (Bou- 
cek 1988). According to Gibson (1989) they 
have either been recorded as primary par- 
asitoids of the fly larvae or as hyperpar- 
asitoids through Platygasteridae (Hyme- 
noptera: Platygasteroidea). The biologies 
of Australian Neaimstatus are largely un- 
known, although one species has been 
reared from galls on Ereinocitrus (Ruta- 
ceae) (Naumann 1991) and CJB has seen a 
specimen reared from an unidentified gall 
on Brachychiton discolor F. Mueller (Steri- 
culiaceae). One Australian species, Neaji- 
astatus cinctiventris Girault, is known to be 
a parasitoid of the Rice gall-midge, Orseo- 
lia oryzae (Wood-Mason), throughout 
southeast Asia. This is the first record of a 
species of Neanastatus attacking a fergu- 
soninid fly. Interestingly, Neanastatus has 
not been reared from several hundred ce- 
cidomyiid galls collected from Melaleuca 
qui}n]uenervia (unpublished data). 

Cinvspilus sp. (Eulophidae). — A single 
species of Cinvspilus was moderately com- 
mon at Peregian where it was the third 
most numerous species emerging from 
galls (Table 1). However, Cinvspilus sp. 
was rare at Morayfleld and was not recov- 
ered from galls at Woodburn. Molecular 
sequences of adults from Morayfleld and 
Peregian were idenflcal (Table 2). In total, 
this species comprised 6.6% of the speci- 
mens reared. Cinvspilus is a large, mor- 
phologically diverse, cosmopolitan genus 



172 



JtlUKNAL OF HyMENOPTERA RESEARCH 



with something in the order of 60 de- 
scribed species from AustraHa (Boucek 
1988). In the absence of any revisionary 
work on Australian Cirwspnhis, it was not 
possible to identify the species from M. 
qui)n]iieuervia galls. However, it belongs to 
a grc^up of species that roughly corre- 
sponds to A. A. Girault's genus Gi/rolnselln 
that was synonymised with Cirrospnlus by 
Boucek (1988). The color pattern of the 
species reared from Fcr^^usonijin sp. galls 
was similar to that of a number of Austra- 
lian Cirrospiliis species that have the body 
mostly yellowish with metallic green 
niarkings on the head and mesosoma and 
a series of dark brown or black transverse 
bands on the gaster. The species in our 
study was similar to the Cirrospiliis reared 
from Fergiisoiiiiwi flaviconiis Malloch galls 
by Taylor et nl. (1996, Fig. 14) but had less 
extensive metallic green on the occiput 
and the median lobe of the mesoscutum. 

Mature larvae of the Cirrospiliis sp. 
reared in this study were distinctive and 
easily distinguished from those of other 
wasps associated with the galls. The larval 
head capsule was virtually prognathous, 
dorsoventrally flattened and with broad, 
cheek-like, lateral expansions basally. The 
mandibles were sickle-shaped and uni- 
dentate. The head, thorax and abdomen 
appeared more or less glabrous, without 
any conspicuous setae. The thorax and ab- 
domen had three and eight low, dorsal 
protuberances respectively. 

The available evidence indicated that 
Cirrospiliis sp. developecl as a solitary, pri- 
mary ectoparasitoid of third instar Fergu- 
sonina larvae. In all instances, Cirrospiliis 
larvae and pupae were found singly, in- 
side isolated, intact chambers. Chambers 
with larvae usually also contained pale 
granules of host remains and Fergiisoiiiiin 
dorsal plates. We also observed intact but 
shrivelled third instar Fcrgusonina larvae 
together with Cirrospiliis larvae. We found 
no evidence of Cirrospiliis larvae feeding 
on gall tissue or acting as hyperparasi- 
toids. On first inspection, chambers with 



Cirrospiliis pupae usually appear empty of 
host remains but contain a thick patch of 
meccinium. On closer inspection, plates 
from Fergiisoiiiim dorsal shields were near- 
ly always incorporated into the patch of 
meconium. 

Cirrospiliis is a biologically diverse ge- 
nus with species developing as parasitoids 
or hyperparasitoids, comn"ionly of leaf- 
miners, or of other larvae and pupae in 
concealed situations (Boucek 1988). In 
Australia, niany species are associated 
with leaf galls, especially on eucalypts 
(Boucek 1988). Taylor ct nl. (1996) reared a 
species of Cirrospiliis from leaf-galls of Fer- 
giisoiiina flaviconiis on Eucalyptus cnnialdu- 
Iciisis. 

Bracoii sp. (Braconidae). — A single spe- 
cies of Bracoii was recovered in relatively 
low numbers from all the sites comprising 
5.8 % of the specimens reared, but it was 
the second most comnion species at 
Woodburn (Table 1). Molecular sequences 
of adults from Woodburn and Peregian 
were identical (Table 2). Bracou is a very 
large, cosmopolitan genus with many 
Australian species, most of them undes- 
cribed (Austin and Faulds 1989, Quicke 
and Ingram 1993). 

Mature larvae of Bracoii sp. were distin- 
guished from those of most other wasps 
associated with the galls, except Poecilo- 
cryptus nigroiuaculaius (see below), by their 
large size. They were also characterized by 
distinctive labial and maxillary sclerites, 
and large, heavily sclerotized, unidentate 
mandibles, which had a series of serra- 
tions on their inner margins. 

Typically Bracou larvae fed indiscrimi- 
nately on hosts within the galls, entering 
multiple chambers and consuming a suc- 
cession of Fcrgusouiua larvae and the lar- 
vae and pupae of the other wasps associ- 
ated with the galls. Often two or more Bra- 
con larvae completed development within 
the same gall. Galls occupied by mature 
Bracou larvae usually had several cham- 
bers interconnected by relatively large 
holes. The chambers were generally 



Volume 10, Number 2, 2001 



173 



packed with brown, particulate debris that 
we concluded was masticated gall tissue. 
Chambers also frequently contained dis- 
sociated Fergiisouiua dorsal-shield plates. 
The remains of a Nenjuistatus pupa and a 
small Poecilocn/p)tiis uigromaculntiis larva 
were also found within chambers occu- 
pied by Bracon larvae. On one occasion a 
Briicoii larva was directly observed feeding 
on a Eiirytoma larva (Table 2). This is the 
first record of a species of Bracon acting as 
a facultative hyperparasitoid. Other 
known species of the genus are primary 
ectoparasitoids (Shaw and Huddleston 
1991). Pupation occured in a relatively 
loosely woven silk cocoon with brown de- 
bris incorporated on its outer surface. The 
cocoon usually filled two gall chambers 
and had a mass of dark meconium depos- 
ited at one end. 

Species of Bracon attack diverse hosts 
but many are parasites of concealed lar- 
vae, mostly of Lepidoptera but also Cole- 
optera and Hymenoptera-Symphyta 
(Quicke and Ingram 1993). Several species 
also parasitize Diptera, particularly gall- 
making larvae (Quicke and Sharkey 1989). 
This is the second record of a species of 
Bracon from a fergusoninid gall, Taylor ct 
al. (1996) having reared B. fergusoniinis 
Taylor, Austin and Davies from Fergnson- 
ina flavicornis leaf-galls on Eucalyptus ca- 
maldulensis. Currie (1937) also reared an 
unidentified braconid from F. nicholsoni 
flower-bud galls on £. niacrorhyncha. He 
reported that the braconid larvae feed "in- 
discriminately on gall tissues and fly lar- 
vae" and it seems likely that their biology 
is similar to the Bracon sp. in our study. 
However, although we confirm that the 
Bracon larvae masticate a considerable 
amount of gall tissue, evidenced by copi- 
ous amounts of brown debris, it is unclear 
whether they derive nutrition from this 
activity or just mechanically scrape away 
the tissue to gain access to additional 
chambers. Larval phytophagy is very rare 
in the Braconidae and has never been con- 



firmed for the subfamily Braconinae (Tay- 
lor ct al. 1996). 

Eupclmus (Macroneura) scniiputata (Gi- 
rault) (Eupelmidae). — Eupelmus scniiputata 
was moderately common from galls at 
Morayfield but rare at Peregian and not 
collected from Woodburn (Table 1). Of the 
103 reared in the study (5.1% of total spec- 
imens reared), 74 came from galls collect- 
ed in 1998. During 1999 only ten E. scnii- 
putata were reared. We did not encounter 
any larvae in our dissections. Molecular 
sequences of adults from Morayfield and 
Peregian were identical (Table 2). There is 
only a single described Australian species 
of Eupclmus {Macroneura) although Boucek 
(1988) indicated a second, presumably un- 
described, Australian species. The speci- 
mens reared in this study appeared to 
match the holotype of £. scniiputata. 

Species of Eupclnius (Macroneura) are 
cosmopolitan, primary or secondary par- 
asites of a wide variety of insect hosts in 
concealed locations, such as within galls, 
grass stems, or cocoons. Some species are 
highly polyphagous, sometimes attacking 
hosts from several different orders (Gib- 
son 1990). A. A. Girault, in his unpub- 
lished manuscript (see Dahms 1978), re- 
corded E. scniiputata emerging from ceci- 
domyiid galls on Pitted bluegrass, Botli- 
riochloa decipicns (Hackel) C. E. Hubbard 
(as Andropwgon pcrtusus (L.) Willdenow). 
Several other species of chalcidoids were 
also reared from these galls. CJB has also 
reared specimens of E. scniiputata from fi- 
nal instar larvae of Aspidoniorpha dciista 
(Fabricius) (Coleoptera: Chrysomelidae), 
most probably as a hyperparasitoid 
through an unidentified tachinid. This is 
the first record of E. scniiputata emerging 
from galls of Fergusoninidae. 

Clironicurytonia sp. (Pteromalidae). — 
Specimens of Clironicurytonia were reared 
in low numbers from all three sites (Table 
1) and comprised 4.6% of the total speci- 
mens reared. Molecular sequences of the 
D2 gene were obtained only from adults 
and larvae from Morayfield (Table 2) but. 



174 



Journal of Hymenoptera Research 



based on morphology, adults from all 
three sites appear to be the one species. 
There are fourteen described species of 
Chroiuciirytomn , all from Australia. The 
species reared in our study could not be 
assigned to one of the described species. 

The larvae of Oiroiuciinjtonia sp. were 
normally active and conspicuously setose. 
They were relatively elongate, gradually 
tapering posteriorly. The body also ta- 
pered anteriorly to a relatively small head 
capsule. The mandibles of mature larvae 
were thin with a single, strong, acute 
tooth. The heavily setose bodies of Chro- 
metin/tomn larvae distinguished them from 
most other larvae within the galls. Ncnii- 
nstntus larvae were superficially sin^iilar 
but less setose, lacking the elongate ven- 
trolateral setae found on the abdominal 
segments of ChwDiciiri/toiim larvae. In ad- 
dition, Chroincunjtoina larvae had lateral 
setae on the first abdoniinal segment (ab- 
sent in Nennnstntiis) and had the most pos- 
terior pair of setae on the head capsule 
more widely separated. The bases of the 
posterior setae on the head capsule were 
separated by more than twice the length 
of a seta in Chwiiiciiri/touin larvae, but only 
by about the length of a seta in Ncniuistatiis 
larvae. Cliwnicuri/toiun larvae usually had 
a conspicuous dorsal hump between the 
first and second abdominal segments and 
a series of thin, transverse, sclerotized, in- 
tersegmental bands between the thoracic 
and first seven abdominal segments. 

Chwiiiciiri/tODin larvae were most com- 
monly solitary hyperparasitoids through 
other Hymenoptera within the galls, feed- 
ing on their mature larvae or pupae. Chro- 
mciii'i/toiiin larvae or pupae were recovered 
from chambers containing the remains of 
Ncanastntus and Eiiri/tonin pupae and from 
chambers containing moribund Brncoii lar- 
vae or their head capsules. On two occa- 
sions, Chrouicuryioma larvae occupied iso- 
lated chambers containing the remains of 
lightly sclerotized Fcrgusouimi larvae, and 
possibly developed as primary ectoparas- 
itoids of the fly. 



Species of Clinviiciinftoiun have been 
reared from unidentified galls on species 
of Eucahjpius and Acacia (Boucek 1988). 
This study is the first to record Oiroinciir- 
i/toiJia emerging from galls of Fergusonin- 
idae and the first to document the larval 
biology of a member of the genus. 

Mcgastig)}ius sp. (Torymidae). — Speci- 
mens of Mcgastigiiius were reared in low 
numbers from all three sites (Table 1) and 
comprised 4.0% of the total specimens. Of 
the 80 adults reared during the entire 
study, 55 emerged from galls collected in 
1997. Molecular sequences of adults from 
Morayfield and Woodburn were identical 
(Table 2). Mcgnstigiiius is a large genus dis- 
tributed throughout most of the world, ex- 
cept the Neotropics. It is particularly spe- 
ciose in Australia with 47 described spe- 
cies (Boucek 1988). In the absence of any 
revisionary studies on the genus and giv- 
en that species often display considerable 
variation in size, color and sculpture (Bou- 
cek 1988), no attempt was made to iden- 
tify the species reared in our study. 

Larvae of Mcgastignius sp. were not en- 
countered in our original dissections of 
galls from which specimens were se- 
quenced. However, examination of larval 
exuviae recovered froni Fcrgusoiiiiia pu- 
paria from which Mcgastigiuiis sp. adults 
had emerged, enabled us to identify lar- 
vae of Mcgastigniiis sp. in subsequent gall 
dissections. Larvae resenibled those of 
Eiiryioina sp., but mature Mcgastigniiis lar- 
vae could be distinguished froni those of 
Eiirytonia sp. and all other gall-associated 
wasps by their distinctive mandibles. Each 
niandible was 4-dentate, with a large, 
acute, apical tooth and three small teeth 
evenly spaced along its inner cutting edge. 
They closely resemble the larval mandi- 
bles of Mcgastigniiis dorsalis (Fabricius) fig- 
ured by Askew (1966). 

The larval biology of Mcgastigniiis sp. 
was variable. Most commonly, adults 
emerged from intact Fcrgusoiiiiia puparia 
found within isolated gall chambers. Each 
parasitized puparium contained the exu- 



Volume 10, Number 2, 2001 



175 



vium of the final instar Mcgnstigiiiiis larva 
and meconium in the form of numerous 
black, discrete pellets. Presumably, Mcgas- 
tigiiiiis sp. developed as a primary para- 
sitoid, either as an endoparasitoid or ec- 
toparasitoid of the pharate Fergusonina 
pupa. We found no evidence that Mcgns- 
tigniiis developed as a hyperparasitoid 
within puparia. Less commonly, Megnstig- 
imis appeared to develop as a solitary ec- 
toparasitoid of third instar Fcrgusoiiiini. 
Megnstignius larvae and naked pupae were 
found within isolated gall chambers that 
contained pale, granulate host remains in- 
cluding dissociated Fergusonina dorsal 
shield plates. In these cases the voided lar- 
val meconium consisted of a thick mass 
instead of discrete pellets. In addition, Me- 
gastigtnus sp. also developed as a hyper- 
parasitoid through Brnco}! sp. On several 
occasions, Megastignnis larvae, pupae or 
pharate adults were found within cocoons 
with the remains of Bmcou prepupae, pu- 
pae or pharate adults. 

Megastignnis is a biologically diverse ge- 
nus with species having larval biologies 
ranging from obligate plant feeders to ob- 
ligate parasitoids (Boucek 1988). Currie 
(1937) reared two species of Megastignnis, 
M. qiiinqiiesetae (Girault) and an unidenti- 
fied species, from Fergusonina nieholsoni 
flower-galls on Eiicalyptiis niacrorln/nelia. 
He reported that the larvae of both species 
were inc^uilines within the galls, forming 
their own separate chambers and presum- 
ably feeding on gall tissue. In contrast, the 
Megastignnis in our study appears to be 
entirely entomophagous. Taylc^r et al. 
(1996) also reared two species of Megastig- 
miis from Fergusonina flavicornis leaf-galls 
on E. eamaldulensis but they did not inves- 
tigate their larval biologies. 

Eupelnius {EupK'hnus) sp. (Eupelmi- 
dae). — Specimens of Eupelmus were recov- 
ered in very low numbers from all three 
sites and accounted for 0.5% of total spec- 
imens reared (Table 1). They appeared to 
belong to a single species and the molec- 
ular sequences of adults from Morayfield 



and Peregian were identical (Table 2). Eu- 
pelnius sp. larvae were not sequenced as 
none were encountered during dissec- 
tions. A single adult female was found in 
an isolated gall chamber together with a 
lightly sclerotized, collapsed Fergusonina 
prepupa. The pupal exuvium of the wasp 
was also present in the chamber. The pre- 
cise larval biology of Eupelmus sp. is un- 
known although it is clearly a solitary pri- 
mary parasitoid or hyperparasitoid. There 
are many species of Eupelmus {Eupelmus) 
found throughout the world; they are par- 
asitic, or rarely 'predatory', on a wide va- 
riety of hosts (Boucek 1988). Harris (1982) 
also reported a species of Eupelmus emerg- 
ing from Fergusonina syzygii Harris galls 
on Syzygium eumini (L.) (Myrtaceae) in In- 
dia. 

Eiuierns sp. (Eulophidaej. — Specimens 
of Euderus were recovered in very small 
numbers from all three sites and account- 
ed for 0.5% of total specimens reared (Ta- 
ble 1). They appeared to represent a single 
species and the molecular secjuences of 
single adults from Peregian and Wood- 
burn were identical. Euderus larvae and 
pupae were not encountered during dis- 
sections. However, the D2 gene sequence 
of a single egg matched that of the adult 
Euderus (Table 2). The egg was attached to 
a moribund Poeeilocrx/ptus nigromaenlatus 
(see below) larva that had been feeding on 
a Fergusonina puparium. Evidently Eude- 
rus sp. acts a hyperparasitoid within the 
galls, which might explain its low relative 
abundance. Euderus is a large cosmopoli- 
tan genus with species attacking larval 
Lepidoptera and Coleoptera (Boucek 
1988). Species are also known to be hy- 
perparasitic, attacking Braconidae (Boucek 
1988). Taylor et al. (1996) also reared a spe- 
cies of Euderus in low numbers from Fer- 
gusonina flavieornis leaf-galls. They sug- 
gested that its larvae might be hyperpar- 
asitic on Bracon fergutoninus within the 
galls. 

Poeeiloeryptus nigromaenlatus Cameron 
(Ichneumonidae). — Poeeiloeryptus nigroma- 



176 



Journal of Hymhnoptera Research 



culatus was the rarest gall-associated wasp 
species at 0.3% of the total specimens 
reared, with only one and four specimens 
recovered from Morayfield and Wood- 
burn respectively (Table 1). The specimens 
appeared to be P. iiigroinaculntiis, although 
they differed slightly in coloration, lacking 
the black markings on the second gastral 
tergite normally found in this species 
(Gauld and Holloway 1986). 

Mature P. uigroiimciilntiis larvae were 
distinguished from most gall-associated 
wasp larvae, except those of Bracou sp., by 
their large size. They could be distin- 
guished from Bracou larvae by their very 
large, heavily sclerotized, bidentate man- 
dibles and by a large sclerotized plate on 
the postlabium (see Short 1978). 

Only a single P. nigroinaciilatus larva 
and a single pupa were recovered from 
dissections of galls from Woodburn, Mo- 
rayfield and Peregian but several larvae 
and prepupae were found in additional 
dissections of galls from Bracken Ridge 
and Coolum (QLD). Only a single mature 
larva, prepupa or pupa of P. uigronmciiln- 
tiis was observed per gall. Each gall had 
its internal structure highly modified, with 
most chanibers breached and intercon- 
nected by relatively large holes. The 
chambers were generally packed with 
brown, particulate debris that we conclud- 
ed was masticated gall tissue. However, 
various chambers also contained dissoci- 
ated Fergiisoiiijia dorsal-shield plates, 
empty Fcrgiisoniiin puparia rent with 
large, ragged holes, the remains of Bracou 
pupae and pharate adults, and Bracou lar- 
val mandibles. Poccilocrypfus ingroiiiacula- 
tus pupated within the gall in a relatively 
large central cavity, incorporating several 
chambers, presumably excavated by the 
larva. Pupation occured inside a brown, 
moderately densely woven cocoon with 
brown debris incorporated on its outer 
surface. 

Poccilocryptus (Subfamily Labeninae) is 
an endemic Australian genus associated 
with a variety of galls on trees of the gen- 



era Eucalyptus, Acacia, Bauksia (Gauld and 
Holloway 1986, Taylor ct al. 1996) and 
now Melaleuca. Poecilocrypius uigroiuacula- 
tus has been reared from anthribid weevil 
galls on Acacia floribuuda (Ventenat) Willd- 
enow { = A. lougifolia), eriococcid galls on 
Eucalyptus (Gauld and Holloway 1986), 
and pteromalid galls on Acacia (Noble 
1941) including those of Trichilogaster aca- 
ciaelougifoliae (Froggatt) on A. florilnuuia, 
and Perilauipella liecataeus (Walker) on A. 
decurreus Willdenow. Poccilocryptus uigro- 
niaculatus has also been recorded from 
galls of Fergusoninidae, Taylor et al. (1996) 
rearing it and P. gallipihagus Gauld and 
Holloway from Fcrguso)^}^ flavicornis 
Malloch galls on Eucalyptus caiiialduleusis. 

There are conflicting reports about the 
biology of Poccilocryptus. We concluded 
that Poccilocryptus iiigroniaculatus larvae 
fed on many hosts within each Fcrgusouiua 
gall, using their large mandibles to chew 
through gall tissue and enter multiple 
chambers. They appeared to consume the 
inhabitants of each chamber regardless of 
its identity. Noble (1941) considered that 
P. Iiigroniaculatus was parasitic upon the 
larvae of a moth that lived as an inc^uiline 
within the multichambered galls of Tri- 
chilogaster acaciaeloiigifoliac. He had no di- 
rect evidence of this host association but 
specifically noted an adult P. nigromacula- 
tus occupying a gall that had much of its 
interior hollowed out by what he assumed 
was a nioth larva. However, his descrip- 
tion bears a striking similarity to the sit- 
uation we observed in the Fergusoiiiiia 
galls and we suggest that P. uigromaculatus 
is probably also a generalist parasitioid 
within Trichilogaster galls. 

Members of the tribe Poecilocryptini all 
appear to oviposit within nutritious plant 
tissue (Gauld and Holloway 1986). The 
enormous mandibles of their larvae led 
Short (1978) to speculate that they may be, 
at least in part, phytophagous. According 
to Quicke (1997) partial phytc^phagy has 
been confirmed for a species of Poccilo- 
cryptus living within coccoid-induced 



Volume 10, Number 1, 2001 



177 



Table 3. Summary of the biologies of Hymenoptera associated with galls of Fergitsonina sp. Eupelmus scm 
ipiitata not included due to lack of information. ? = Some evidence but not confirmed. 



Spe 



iVimai y 
indiipar.isitoid 



Primarv 
r.ctopaiMsitoid 



Hvperp.ird^jtoid 



Mullipk' hosts 



Euri/toma sp. 
Coelocyba sp. 
Neanastatiis sp. 
Cirrospilus sp. 
Brncou sp. 
CJironieiin/toina sp. 
Mcgasti;^nius sp. 
Eiipcliinis {Eiipclniiis) sp. 
Euderus sp. 
Pflccilocryptiis )iigroiimciilatiis 



X 
X 




X 
X 


X 




X 


X 




X 


X 


X 


rarely 


? 


X 


X 


X 


X 


X 
X 




X 




X 


X 





galls. We confirmed that Poccilocrypiiis lar- 
vae masticate a considerable amount of 
Melaleuca gall tissue, evidenced by copi- 
ous amounts of brown debris. However, it 
is unclear whether they derived nutrition 
from the plant tissue or just niechanically 
eroded the chamber walls to gain access 
to their contents. 

CONCLUSION 

Of the eleven species c^f Hymenoptera 
associated with the Fergiisouiua sp. gall, 
we were able to observe and describe the 
biology of nine of the species (Table 3). 
Seven commonly or exclusively fed di- 
rectly upon the Fergusoriiiin larvae and or 
pupae with most species developing on a 
single host. However, three of the seven 
usually fed on multiple hosts. These spe- 
cies have specialized biologies, which en- 
able them to chew through plant tissues 
to access gall inhabitants. On the available 
evidence, the remaining two species ap- 
peared to be hyperparasitoids. We do not 
know which of the eleven species are gall 
specific, but did find it interesting that 
none of these species were reared from 
other galls on M. quinquenervia formed by 
Cecidomyiidae or Homoptera (unpub- 
lished data). None of the Hymenoptera in 
this study could be described as inqui- 
lines. The term inquiline is defined by Tor- 
re-Bueno (1989) as a commensal that lives 
in a very close spatial relationship with its 



host, in its shelter, not feeding on it, but 
frequently destroying it. We did occasion- 
ally observe species of Lepidoptera acting 
as inquilines, including one species, Holo- 
cola sp. (Tortricidae), which is known to 
feed on young leaf and flower buds of M. 
quinquenervia (unpublished data). 

We found the D2 molecular method to 
be robust for characterizing all life stages 
including eggs and small larvae. Further, 
the D2 gene sequence was consistent for 
each species and between adults and im- 
matures. In the dissections we encoun- 
tered the immature forms of eight out of 
eleven species, which were matched with 
the adult forms. The molecular technology 
provides many advantages in the study of 
cryptic immature insects. The amount of 
time and effort required to identify im- 
matures is greatly reduced because rear- 
ing to the adult stage is not needed. The 
biology of the immatures can be observed 
//; vivo and matched with adults without 
speculation or comparison to known bi- 
ologies of related species. A greater num- 
ber of gall-inhabiting insects are likely to 
be discovered using this technique as 
compared to other techniques. Removal of 
the gall from the plant and holding it for 
insect emergence subjects the inhabitants 
to changes in plant turgor, humidity and 
temperature. All of ti^iese factors could be 
critical to the survival of the immatures. 
Sleeving the gall for collection of emerging 



178 



Journal of Hymenoptera Research 



insects may be biased against late arrivals 
such as hvperparasitoids. 

Gene sequences are quantitative and 
can be compared to sequences collected 
later in the study or by other researchers. 
By posting the sequence on GenBank*' 
other researchers may in turn match the 
identity of their insects. Sequence data 
serves as an interim identity for the insect 
species until they are described. Field 
studies and biological control programs in 
particular should submit vouchers not 
only of the insect specimens but also of the 
gene sequences as well. Later revisions of 
genera could include, where possible, the 
molecular data from a wide array of bio- 
logical studies. In this way a greater num- 
ber of specimens could be identified si- 
multaneously. Our understanding of the 
biology and distribution of insect species 
would be greatly enhanced. 

In biological control programs directed 
against weeds, agents must reach high 
population levels in order to control their 
host. Development of high population lev- 
els in the region of introduction is pro- 
moted initially by an almost unlimited 
food supply and by release from the 
agent's natural enemies (Harley and For- 
no 1992). Fergusoniuil sp. is likely to be in- 
troduced to Florida, USA, where it will 
find an abundance of suitable M. quiuque- 
nervia plant bucis which it needs to form 
galls (Goolsby ct al. 2000a, Van ei nl. 2000). 
In its region of origin Fcrgitsouiim sp. is 
heavily attacked by natural enemies, in- 
cluding eight primary parasitoids. One 
would predict that fewer parasitoid spe- 
cies would attack Fcrgusoiiiiin sp. in Flori- 
da, and that they would be less co-adapt- 
ed than those in Australia. Fergusoninidae 
are not represented in the New World, so 
the association with this family of gall- 
making flies would be novel fc^r the indig- 
enous parasitoids. In the absence of its co- 
evolved natural enemies, Fcrgiisoniiia sp. 
could reach much higher populations lev- 
els, potentially having an impact on M. 
qiiinquencrvin. We hope that our study 



provides the basis for future comparisons 
of natural enemies of Fergusoiiiiia sp. in 
both its native and adventitious range. 
This research would further our ability to 
predict the impact of indigenous parasit- 
oids on introduced biological control 
agents. 

ACKNOWLEDGEMENTS 

The authors wish to thank John LaSalle, Mike Rose, 
and the referees for their helpful comments in prep- 
aration of the manuscript; Diana Hartley and Leslie 
McKenzie for molecular sequencing; Matthew Pur- 
cell, Ted Center and Gary Buckingham for collabo- 
rative insights; Jeff Wright for photographs of the 
galls. Research funding was provided by South Flor- 
ida Water Management District, Army Corps of En- 
gineers, Florida Department of the Environment, and 
USDA-ARS-Office of International Research Pro- 
grams. 

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J. HYM. RES. 
Vol. 10(2), 2001, pp. 181-230 

Taxonomy and Ecology of Costa Rican Euplectrus (Hymenoptera: 
Eulophidae), Parasitoids of Caterpillars (Lepidoptera) 

Michael E. Schaufe and Daniel H. Janzen 

(MES) Systematic Entomology Laboratory, Agricultural Research Service, Plant Sciences 

Institute, c/o National Museum of Natural History, Washington, D.C. 20560-0168, USA (e-mail: 

mschauff@sel.barc.usda.gov); (DHJ) Department of Biology, University of Pennsylvania, 

Philadelphia, PA 19104, USA (e-mail: djanzen@sas.upenn.edu) 



Abstract. — The species of parasitic wasps in the genus Euplectrus (Hymenoptera: Eulophidae) 
known from Costa Rica are reviewed, and their ecology is briefly summarized from a long-term 
on-going in\'entory of caterpillars (Lepidoptera) and their parasitoids in the Area de Conservacion 
Guanacaste in northwestern Costa Rica. Twenty species are reported, of which 17 are new species 
described by Schauff: Euplectrus anae, E. carloivae, E. edithae, E. floryae, E. hansoni, E. ireneae, 
E. ivonae, E. josei, E. magdae, E. mariae, E. orias, E. rojasi, E. ronniei, E. imlvcrdei, E. ivalteri, 
E. xiomarae and E. znmorai. An illustrated key and photographs of larvae supplement the de- 
scriptions. 



Members of the genus Euplectrus (Hy- 
menoptera: Eulophidae) are parasitoids on 
many species of caterpillars that live and 
feed exposed on the foliage of their food 
plants (Ferriere 1941, Boucek 1988). Some 
species have been used in biological con- 
trol (Puttier ct al. 1980). The wasps' ability 
to arrest host caterpillar molting by inject- 
ing a chemical arrestant through the ovi- 
positor prior to egg laying also has poten- 
tial as a tool in pest control (Coudron and 
Puttier 1988, Coudron and Brandt 1996). 
In spite of its potential value to agriculture 
and intriguing natural history, Euplectrus 
has not been studied in any comprehen- 
sive manner. This is the first in a series of 
studies anticipated on the systematics of 
the New World species of Euplectrus. It is 
intended to alert field biologists about 
their distinctive ecology. 

Very little is known of the systematics 
of the Central American species of this ge- 
nus, and almost nothing has been pub- 
lished on their ecology/ life history. An 
unpublished Ph.D. thesis discussed the 
New World species and contains records 
of several Central American species (Gon- 



zalez 1985). But, since this thesis has re- 
mained unpublished, these records re- 
main unavailable. DeSantis (1967), De- 
Santis (1979), and DeSantis (1980), De- 
Santis and Fidalgo (1994) cataloged the 21 
species that have been described from 
Central and South America. Only three 
species were recorded from Costa Rica (£. 
cosintockii, E. furuius, and E. solitarius) and 
we describe 17 new ones in the paper. 

As is often the case in chalcidoid wasps, 
host records for Euplectrus have been scat- 
tered and are of cquestionable accuracy. 
However, extensive rearing of Lepidop- 
tera larvae for the past two decades (http: 
//janzen. sas.upenn.edu) as a caterpillar 
inventory of the biodiversity of the Area 
de Conserxacion Guanacaste (ACG) in 
northwestern Costa Rica (http: //www. 
acguanacaste.ac.cr and Janzen 2000) has 
generated more than 250 Euplectrus rear- 
ings, along with those of other parasitoids 
(e.g., Gauld et al. 1992, jcinzen 1993, Gauld 
and Janzen 1994, Sharke)' and Janzen 
1995, Dangerfield et al. 1996, Janzen, D. H. 
and I. D. Gauld 1997, Zitani et al. 1997, 
Janzen et al. 1998). In addition, P. Hanson 



182 



Journal of Hmvienoptera Research 



of the Universidad de Costa Rica and I. D. 
Gauld of The Natural History Museuni 
have extensively malaise trapped the par- 
asitoid fauna in Costa Rica, and this ma- 
terial is considered here as well. As a re- 
sult of these efforts, we are now able to 
report in a detailed manner on the com- 
position of this genus over a large area of 
many habitats. Although this study ex- 
pands the number of species known from 
Costa Rica by about 500%, we feel that 
many more species remain to be discov- 
ered. Eiiplcctnis appear in Malaise traps 
with very low frequency, so that their 
presence will have to be detected largely 
through rearing programs. 

Among Eulophidae, the tribe Euplectri- 
ni is one of the most easily identifiable 
since all species share the greatly elongat- 
ed hind tibial spur(s) (Fig. 89) that have 
been the defining synapomorphy for the 
group since the last century (Boucek 1988, 
Wijesekara and Schauff 1994). Species of 
Euplectrus can be differentiated from oth- 
er genera in the Euplectrini by having a 
simple median carina on the propodeum, 
lacking subniarginal grooves on the scu- 
tellum, and having only 2-3 pairs of setae 
on the mesoscutal midlobe and spinning a 
cocoon as illustrated in Figs. 1-9. A key to 
New World genera was published by Wi- 
jesekara and Schauff (1997). 

Euplectrus (Fig. 4) is also unique among 
Eulophidae in that the larvae live exter- 
nally on the host and spin a cocoon in 
which to pupate. Equally unique, the co- 
coon silk is produced from the anus by 
modified malpighian tubules (Ferriere 
1941). The eggs are laid externally in 
groups on the host caterpillar. The larvae 
feed on hemolymph through the cuticle 
and mature while attached to the back of 
the host by their mouthparts (Figs. 1, 5, 6, 
8). When ready to pupate, the larvae of 
son^ie species move to the underside of the 
caterpillar cadaver (Figs 2, 3, 9), while oth- 
ers spin a ruff of cocoons around the ca- 
daver. 

Below, we describe the species of Eu- 



plectrus (the senior author is the taxonomic 
author of these species), and give an ac- 
count of the natural history for those on 
which we have been able to accumulate 
data. 

Abbreviations of museums are as fol- 
lows: U.S. National Museum of Natural 
History, Washington, D. C. (USNM); The 
Natural History Museum, London 
(BMNH); Instituto Nacional de Biodiver- 
sidad (INBIO), Costa Rica; Canadian Na- 
tional Collection, Ottawa (CNC). 

High resolution digital copies of the 
original photographs for Figures 1-9, all 
taken in the Area de Conservacion Guan- 
acaste, are available at Janzen and Hall- 
wachs (2000). 

Morphology . — One of the distinctive fea- 
tures of species of Euplectrus is the ar- 
rangement of small and enlarged setae on 
the head (Fig. 23). There are up to six pairs 
of thickened and elongated setae on the 
upper part of the head. There are two 
pairs postero-laterally along the occipital 
carina (SI and S2). There is a pair between 
the posterior ocelli (S3) and a pair laterad 
of the anterior ocellus (S4). S5 lies on the 
vertex forward of S4 and between the top 
of the scrobes and the eye. S6 is located 
near the edge of the eye about 1/2 way 
down the orbit. In all the species exam- 
ined to date, SI, S2, S3, and S6 are always 
present. S4 and S5 may be either reduced 
or absent. 

In addition to the enlarged setae, there 
are also several smaller setae, some of 
which occur in pairs (Fig. 23). There is 
generally a row of setae behind the eye 
and below the occipital carina (srl). There 
is a pair of small setae (ssl) near the oc- 
cipital carina between SI and S2. Between 
and slightly behind the posterior ocelli are 
one or two pairs of small setae (ss2) that 
are occasic^nally absent. Laterad of S4 
there is a single small seta (ss3) and lat- 
erad and below S6 there is an irregular 
line of setae (sr2) that usually extends to 
the bottom of the eye. 

The antenna has four funicular seg- 



Volume 10, Number 2, 2001 



183 








Figs. 1-3. Lepidopteran larvae attacked by Eiiplcctiiis. I, A normal size clutch ul L. flon/ae larvae feeding on 
body fluids of a pen-penultimate (third) instar of Ein/o oci/petc (Sphingidae). All species of Euplcctrus for which 
the larvae are known except £. waltcri (Figs. 4-7) display this general appearance on their host caterpillars 
(e.g., Fig 8); 2, The mummified cadaver of an ultimate (fifth) instar Pncctcs limodcs (Noctuidae) caterpillar after 
the larvae of E. joesi have terminated their feeding on it and spun cocoons between the cadaver and the leaf, 
firmly silking/gluing the cadaver to the cocoons (see Fig. 3). Fourteen spheroidal meconial pellets from 14 
wasp larvae surround the cada\'er. This is the pupation method used by all species of Euplcctrus kno\vn except 
for E. waltcri (Figs. 4-7); 3, The side-by-side and often head-to-toe cocoons/pupae of 16 £. josci after the 
mummified cadaver of their host caterpillar (and see Fig. 2) has been stripped away. When exposed in this 
manner, the pupae often die of apparent desiccation unless placed in a high-humidit\ container. 



ments (F1-F4) followed by a club or clava in females) to greatly enlarged and bal- 

(Fig. 25). The female scape is cylindrical loon-like (Fig. 50). a\11 male scapes possess 

and 4-6X as long as wide. The male scape groups of sensillae which appear as round 

shows great variation from cylindrical (as clear areas in slide-mounted specimens 



184 



Journal of Hymenoptera Research I 




Figs. 4-6. SphingiJ Ijiaju jtUickcJ b\ L(//'/(l ////.-. uuillcn. 4, \\\o tcnialcs waiting to oviposit alter this pen- 
penultimate (third) instar Maiidiica bnnicsi has molted to penultimate instar, the instar usually parasitized by 
Eiiplixtnis walteri when attacking the larger of the species of Manduca. This caterpillar was within 24 hours of 
molting. The wasps clung tenaciously to the caterpillar despite being banged around in a plastic bag and 
roughly transported from the forest to the laboratory where the photograph was taken. 5, One or several 
clutches of partly grown Euplcctnis walteri larvae feeding on an ultimate instar Maudiicn dilucida (dark morph). 
6, A clutch of full-grown larvae just beginning to spin the first strands of silk of their aggregate cocoon on 
the back of a just-died penultimate (fourth) instar Manduca floivstaii. The distended ceratopogonid fly sucking 
caterpillar blood on the far right is Forciponn/in prob. fuli^^iiiosa (Art Borkent, pers. comm.). 



Volume 10, Number 2, 2001 



185 



(Figs. 19-21). The patterns of these sensil- 
lae along with the shape of the scape is 
often species-specific (Figs. 27-39). 

Abbreviations for measurements and 
ratios are as follows (Figs. 23, 24): OD: 
OOD is the longest ocellar diameter com- 
pared to the distance between the ocellus 
and the eye (ocell-ocular distance); FW: 
EW is the ratio of the width of the face 
(distance between the margins of the eyes 
at the midpoint) to the width of the eye 
(distance from the margin of the eye to the 
edge of the eye when viewed from the 
front of the head). Height of the eye (EH) 
is the distance from the bottom of the eye 
to the top of the eye when viewed either 
from the front or laterally. The malar 
space (MS) is the distance from the bottom 
of the eye to the oral opening. 

Rearing. — Wild-caught caterpillars were 
individually reared and databased as part 
of an ongoing inventory (http://janzen. 
sas.upenn.edu and Janzen and Hallwachs 
2000) of all the caterpillars of the dry for- 
est, cloud forest, and rain forest of the 
Area de Conservacion Guanacaste (ACG) 
in northwestern Costa Rica (http:// 
www.acguanacaste.ac.cr). Each caterpillar 
was maintained in a plastic bag or glass 
bottle at ambient temperatures while it 
and /or its parasitoids developed. When 
parasitoids appeared, or in this case, when 
the Eiiplcctriis larvae were noted on the 
caterpillar, the caterpillar was isolated in 
a glass bottle with a small amount of food 
plant as a perch, to maintain humidity and 
serve as a substrate for the cocoon-spin- 
ning of the Eupkctrus larvae. Eupkcinis co- 
coons were monitored daily, and when 
the wasps eclosed, they were preserved in 
alcohol for later shipment to the senior au- 
thor. Eupkctrus adults were noteworthy 
for generally living at least 48 hours in the 
rearing container before dying (of starva- 
tion?) while other similarly-sized Eulophi- 
dae, Braconidae and Ichneumonidae rare- 
ly survive more than 24 hours in the rear- 
ing container without being fed. 

This methodology of caterpillar and 



parasitoid surveying suffers the limitation 
that once the caterpillar has been found in 
nature and brought into captivity, it and 
its parasitoids are no longer available for 
parasitization or hyperparasitization. This 
means that the percentages of parasitiza- 
tion recorded here represent the minimal 
possible. Equally confounding is that cat- 
erpillars are captured at all stages in their 
development, with the result that a simple 
comparison of the number of caterpillars 
parasitized against the number of cater- 
pillars captured may severely underesti- 
mate the intensity of parasitization. This 
underestimation is because in the case of 
some species of Euplcctrus, the wasp larvae 
kill the caterpillars during instars prior to 
the last ins tar. The number of "unparasit- 
ized" later instars found is therefore mean- 
ingless vis-a-vis Eupkctrus percent parasit- 
ization. It should be emphasized that all 
sphingid caterpillars that have been found 
in the wild in the inventory are captured 
and reared in captivity, so there is no bias 
generated through seeking just those in- 
dividuals parasitized by Eupkctrus. 

There were two kinds of Eupkctrus iden- 
tifications produced in this study. The most 
certain are those where wasps eclosed. All 
of those were identified by the senior au- 
thor, a eulophid taxonomist. In 20-30% of 
the cases of parasitization by the uniquely 
distinctive external green larvae of Eupkc- 
trus (Fig. 1), the wasp larvae died when at- 
tempting their relocation below the cadav- 
er or died because of severe rearing con- 
ditions (excessive heat, moisttire, desicca- 
tion). For Eupkctrus floryae, E. wnltcri, E. 
mariac and £. josei, the junior author (a Eu- 
pkctrus ecologist) identified those dead lar- 
vae based on their distinctive cocoons, tim- 
ing of attack, and taxon of host. These iden- 
tifications are pooled with those of the se- 
nior author when attempting to tease out 
the intensity of attack and certain questions 
of parasitoid host specificity, but may also 
be treated as two separate classes of data 
at the wishes of the reader, since each re- 
cord is individually documented in the da- 



186 



Journal of Hymenoptera Research 



tabase and the identifier identified (http:/ 
/janzen. sas.upenn.edu). 

The basic collection locality and ciate in- 
formation for all the reared Euplectrus de- 



scribed here (Table 1) is deliberately min- 
imal. More detailed information on a par- 
ticular rearing may be found at http:// 
janzen.sas.upenn.edu. 



Table 1. Host associations for the 11 new species of reared Euplectrus described here from the ACG. All 
records are from wild-caught caterpillars of the species indicated. Details may be found in the individual 
records in Janzen and Hallwachs (2000). 



Euplectrus 
species 



Number i 

lu.sts 
p.ir,isiti/i' 



Number 

u^ceptible 

reared 



anae 
florync 



ircneae 
ivonae 
josci 
uia^dac 



arms 
nviuici 

umlteri 



Spliacclodcs inihierana (Hubner) (Geometridae) 
Enyo ocypcte (L.) (Sphingidae) 
Pcrigoiiia ihis Boisduval (Sphingidae) 
Pcrigoiiia lusca (F.) (Sphingidae) 
AcUopos fadus (Cramer) (Sphingidae) 
Moti/a abscuznlis (Walker) (Noctuidae) 
Euscirrlioptcrus pocyi Grote (Noctuidae) 
Pncctcf luiiodcs (Guent§e) (Noctuidae) 
Dnsylopliia iiiaxtln (Shaus) (Notodontidae) 
Dasylop^hia hasitmcta Dognin (Notodontidae) 
Dasyloplun iir. goraxa Schaus (Notodontidae) 
Coucaiia iiiuiidissinia Walker (Noctuidae) 
Eli/uiiotis attcuuata (Walker) (Notodontidae) 
Dasi/lopliia ur. goraxa Schaus (Notodontidae) 
Geometridae 

Oxidcrcia toxcn (Stoll) (Noctuidae) 
Cautctliia spuria (Boisduval) (Sphingidae) 
MiiiidtiCii bnnicsi (Clark) (Sphingidae) 
Maiiduca dilucida (Edwards) (Sphingidae) 
Manduca florestan (Cramer) (Sphingiciae) 
Maiiduca lanuginosa (Edwards) (Sphingidae) 
Manduca rustica (F.) (Sphingidae) 
Perigonia ilus Boisduval (Sphingidae) 
Hcniiceras clarki (Notodontidae) Schaus 
Hcmiceras corcnia Schaus (Notodontidae) 
Hcniiceras nigrcsccns Schaus (Notodontidae) 
Roscnia attcuuata (Notodontidae) 



1 


197 


0.5 


84 


1010 


8.3 


17 


256 


6.6 


2 


73 


2.7 


4 


256 


1.6 


1 


29 


3.5 


1 


174 


0.6 


11 


73 


15.0 


1 


29 


3.1 


1 


65 


1.3 


1 


12 


8.3 


30 


194 


15.5 


7 


213 


3.3 


1 


12 


8.3 


1 


1885 


0.05 


1 


91 


1.2 


1 


110 


0.9 


1 


34 


2.9 


4 


112 


3.6 


19 


163 


11.7 


3 


68 


4.4 


3 


26 


11,5 


1 


181 


0.5 


6 


183 


3.3 


1 


71 


1.4 


1 


521 


0.2 


14 


111 


12.6 



KEY TO SPECIES OF COSTA RICAN EUPLECTRUS 

1. Either face beneath toruli marked light brown or black (Fig. 15) or hind coxae brown to black 
(Fig. 11) ^. ^ 2 

- Face beneath toruli all yellow (Figs. 12, 13, 16) and hind coxae light yellow or white (Fig. 10) 

9 

2. Apical flagellomeres dark and contrasting with yellow to light brown El (Fig. 22) 3 

- Apical tlagellomeres same color as El or only slightly darker 4 

3. Width of face about 4X width of eye viewed from front (to measure EW:EW, see Fig. 24); 
posterior ocellus about 2X its own diameter removed from the margin of the eye (Fig. 49); 
mandibles yellow, male scapes greatly swollen (Fig. 50) furnitis Walker 

- Width of face about 2.5X width of eye viewed from front; posterior ocellus about 1.25X or less 
its own diameter removed from margin of eye; mandibles brown, male unknown 

zmnorai, new species 



Volume 10, Number 2, 2001 




T^^^f^^^'^'^ 







4^ 



Figs. 7-9. Lepidopteran larvae attacked by Eiiplcctnis. 7, The aggregate cocoon of a clutch of 100-200 Eu- 
p^Icctni^ ivaUcri pupae ringing the semi-decomposed mummy of an ultimate (fifth) instar Maininca dilucidn that 
was their host. The ruff of cocoons is a single dense unit with the cocoons tightly packed together and bound 
together by silk and glue. This is the only species of Eiipilcctnis known to make an aggregate cocoon of this 
nature. 8, A normal-sized clutch of E. xioiiiarac larvae feeding on body fluids of a penultimate (fourth) instar 
Hemiceras darki (Notodontidae). 9, The mummified cadaver of a penultimate (fourth) instar liciuiccras clnrki 
(Notodontidae) caterpillar after the larvae of £. xioiuarac have terminated their feeding on it and spun cocoons 
between the cadaver and the leaf, firmly silking/glueing the cadaver to the cocoons. Three beige spheroidal 
meconial pellets are suspended on the outer surface of the silk aggregate cocoons. This i.-> the pupation method 
used by all species of Euplcctrns known except for £. imltcri (Figs. 4-7). 



188 Journal of Hymenoptera Research 

4. Hind coxa yellow or very light brown (Fig. 10) 5 

- Hind coxa dark brown or black (Fig. 11) 6 

5. First tarsomere of hind leg about ecjual in length to second; postero-lateral margin of scutellum 
overlapping metanotum (Fig. 43) carlowne, new species 

- First tarsomere of hind leg much longer than second; postero-lateral margin of scutellum not 
overlapping metanotum edithae, new species 

6. Posterior margin of scutellum overlapping anterior edge of metanotum medially (Fig. 68, 69); 
anterior median propodeal carina split into a V-shape, not enlarged and cup-like (Fig. 69) . . 

rojasi, new species 

- Posterior margin of scutellum not overlapping anterior edge of metanotum medially, at least 
two distinct alveoli visible at anterior edge of metanotum (Figs. 53, 59); anterior median pro- 
podeal carina expanded and protruding from surface of propodeum, invaginated and cup-like 
(Figs. 42, 43) 7 

7. One pair of small setae (ss2) present between lateral ocelli (Figs. 66, 82) . . orias, new species 

- No small setae (ss2) between lateral ocelli (Fig. 85) 8 

8. Mandibles brown; malar suture present at least below eye margin (as in Fig. 67) 

valverdi, new species 

- Mandibles yellow; malar suture absent xiomarae, new species 

9. Occiput between posterior ocelli with 2 pairs (4) of small setae (ss2) (Fig. 46) 10 

- Occiput between posterior ocelli with 1 pair (2) or no small setae (ss2) (Fig. 49) 11 

10. Yellow color on face extending about 1/2 way up eye margin and above level of toruli (Fig. 
12) flonjac, new species 

- Yellow color on face restricted to below and between toruli, not extending up along margin 
of eye (as in Figs. 13, 14) josei, new species 

11. Posterior ocellus less than 1 diameter from edge of eye (see Fig. 23); width of eye more than 

1/2 width of face (Fig. 24) ireueae, new species 

Posterior ocellus more than 1 diameter from edge of eye; width of eye less than 1/2 width of 
face 12 

12. Toruli separated by about 4X their own diameter (Fig. 13) zimlteri, new species 

Toruli separated by about 2-2. 5X their own diameter 13 

13. Funicular segments Fl-4 all about 3X as long as wide and Fl-3 all about same length as club 

(Fig. 26) haiisoni, new species 

At least one of funicular segments Fl-4 less than 3X as long as wide, generally only 2X as 
long as wide or less; F2-4 usually much shorter than club 14 

14. Enlarged seta number 5 (S5, Fig. 23) present 15 

Enlarged setae number 5 absent 18 

15. Anterior metanotum with a single large alveolus, sometimes divided medially into two large 
alveoli (Fig. 59); propodeum distinctly reticulate ewer entire median surface 

iiiadgne, new species 

- Anterior metanotum with a thin line of small alveoli (Fig. 40); propodeum not distinctly 
reticulate over surface, usually lightly reticulate or smooth and shiny 16 

16 Petiole distinctly wider than long; propodeum adjacent to median carina with irregular small 

carinae, appearing rugose; metasoma light brown to brown in apical half 

anac, new species 

Petiole as long as wide or kmger than wide; propodeum adjacent to median carina smooth 
or very lightly reticulate; metasoma dark brown to black apically 17 

17. Fl about 3X as long as wide solitarius Ashmead 

- Fl about 2X as long as wide comstockii Howard 

18. Yellow coloration of face extending laterally beyond the outer edge of the toruli, often reaching 

to under the eye (Fig. 16) ivoiiac, new species 

Yellow coloration of face restricted to between and below the toruli, occasionally slightly 
laterad of the toruli, but not to near or under eye 19 

19. Propodeum immediately laterad of median carina with small irregular carinae (appearing 



Volume 10, Number 2, 2001 



189 



somewhat rugose) or lightly reticulate; petiole wider than long. Male scape with small brown 
sensory area on ventral edge which extends at most (2/3 or more) of length of scape (Fig. 30) 

with 2-3 irregular rows of sensillae mariae, new species 

Propodeum laterad of median carina mostly smooth; petiole as wide as long. Male scape with 
small brown sensory area on ventral edge which extends for less than 1/2 length with only 
a single row of sensillae (Fig. 31) ronniei, new species 



ECOLOGY 

Eiiplccfnis floryac Schauff. — This species 
is a common parasitoid (Table 1) of sec- 
ond instar Emjo ocypcte (Sphingidae), a 
medium-sized caterpillar that feeds on 
Dilleniaceae in the dry forests of the ACG 
(Fig. 3). It is likely that this small sphingid 
supports most of the Eiiplcctnis floryac 
population in these forests. This Eiiplcctrus 
has not been found in any of the ACG rain 
forest or cloud forest. Parasitized Euyo 
oci/petc larvae have been found feeding on 
Tctrnccrn volubilis L. and CuratcUa aiiicri- 
cniin L. This sphingid feeds also on Dnvilhi 
kufitfiii A. St.-Hil. and Doliocnrpus dciitatiis 
but in such low numbers that the absence 
of Eiiplcctrus florync records on these cat- 
erpillar food plants may be a sampling ar- 
tifact. The parasitized Euyo ocypcte cater- 
pillars were found in sites ranging from 
fully isolated habitats (mostly CiirntcUii 
amcricana) to the deep shade of old-growth 
forest (mostly Tctrnccrn volubilis L.). 

Only one Enyo ocyf>ctc wild-caught first 
instar was found to have Eiiplcctrus floryac 
on it, and that one died without its three 
parasite larvae completing development 
(92-SRNP-835). However, many Euyo ocy- 
pcte larvae have been found in nature as 
second and third instars with Eiiplcctrus 
floryac feeding on them. Only two fourth 
and no fifth (last) instar Euyo ocypcte lar- 
vae have been found with Eiiplcctrus flor- 
yac on them. These records imply that 
once the caterpillar has reached the pen- 
ultimate or last instar, it is imniune or un- 
attractive to the wasp adult. It appears 
that oviposition normally occurs on sec- 
ond or third instar larvae. A single cater- 
pillar normally supports 5-10 Eiiplcctrus 
floryae larvae in a single group. We as- 



sume that this represents a single ovipo- 
sition by a single female. 

There is no suggestion that the sphingid 
caterpillar molts after wasp oviposition 
occurs. The wasp larvae develop from mi- 
nute, glabrous, green protuberances in a 
cluster to a tight cluster of large globular 
green larvae (Fig. 3) at the same point on 
the back over a period of 4-8 days. They 
do not move about and feed with their 
heads always inserted in the same hole. 
The caterpillar does continue to eat leaves 
during this period, but stops feeding in 1- 
2 days before the wasp larvae release their 
feeding position and move to underneath 
the moribund caterpillar to spin their co- 
coons on the leaf underside (e.g. Figs. 2, 
3) (where the caterpillar also rests when 
unparasitized). The beige/brown silk co- 
coons are adjacent with their long axis at 
right angles to the long axis of the cater- 
pillar, alternately c^riented to the left and 
right, with the meconial pellet clearly vis- 
ible at the tip of the cocoon. The cocoons 
are tightly glued between the ventral sur- 
face of the cadaver anci the leaf. The co- 
coons are enmeshed in a distinctive silken 
matrix resembling a loose basket. The ca- 
daver becomes a mummified strap of tis- 
sue perched on top of the cocoons. 

The wasps spend 7-12 days in their co- 
coons. However, while wasp eclosion date 
is accurately recorded (see text account of 
specimens examined), some of the shorter 
development times of 7 days may be cases 
where the cocoons were not noticed for 1- 
2 days after they were spun. It is likely 
that the usual time in the cocoon is about 
8 days. There is no evidence of dormancy 
by pupae in cocoons spun at any time in 
the rainy season. Only once has a cater- 
pillar parasitized bv Eiiplcctrus floryac been 



190 



Journal of Hymenoptera Research 



found in the dry season, and in this case 
there was no indication of dry season dor- 
mancy (92-SRNP-324). 

The overall phenology of Ein/o oci/pctc 
second and third instars in this dry forest 
habitat is that they first appear on Ciirii- 
tella nniericniin L. in small numbers in Feb- 
ruary-April, and then in much larger 
numbers in May-June. In June-July, there 
are large numbers on Tetracem voliibilis L., 
followed by a very few individuals on 
both species of plants through December. 
One Euplectrus flori/ae record is in Febru- 
ary and one at the end of September. As 
a general pattern, it appears that the Eu- 
plectrus floryae population could have as 
many as 4-5 consecutive generations on 
the Enyo ocypclc population beginning in 
early May-July. It survives the August-to- 
April last half of the rainy season and 
nearly all of the dry season as non-repro- 
ducing adults. It is unknown whether they 
migrate to a wetter part of Costa Rica, or 
"hide" in local moist areas within the dry 
forest. They do not appear in Malaise 
traps at any time of the year, even when 
these traps are in the middle of the forest 
habitats in which they are breeding, only 
a few meters from parasitized caterpillars. 
The overall results of the caterpillar rear- 
ing program in this forest suggests that 
Euplectrus floryae does not use an alternate 
host caterpillar species or family at these 
other times of the year. 

The few records of Euplectrus floryae 
from Cautetliia spuria, two records from 
Aellopos fadus, and 18 records from Peri- 
goiiia ilus and Perigouia lusca (these two 
species are nearly indistinguishable as lar- 
vae) (Table 1), probably represent minor- 
ity hosts for Euplectrus floryae. These four 
sphingids suffer no more than 27o parasit- 
ization by E. floryae, while, for example, of 
200 larvae of Eiiyo ocifpete collected in the 
second and third instars (the key suscep- 
tible stages), 25% had Euplectrus floryae on 
their backs. The parasitization of non-£//i/o 
ocypete caterpillars occurs at the same time 
of year (the first half of the rainy season) 



as do the bulk of the rearing records from 
Euyo ocypete. 

Euplectrus floryae is essentially the only 
hymenopterous parasite in the ACG dry 
forest habitat that kills Einjo ocypete prior 
to the last instar (except for two braconid 
rearings and one ichneumonid from 1240 
caterpillars). The other parasitoids of Enyo 
ocypete are two species of Tachinidae; their 
larvae emerge from the caterpillar at the 
end of the last instar {Driuo piceiventris) or 
the adult fly ecloses from the pupa {Bel- 
vosia sp.). While this forest has thousands 
of species of hymenopterous and dipter- 
ous parasitoids of caterpillars, Euplectrus 
floryae shares Enyo ocypete with only two 
of them. The two fly parasites may get 
into their hosts as early as the second in- 
star, but more commonly do so in the 3- 
5th instars. In effect, Euplectrus floryae uti- 
lizes its portion of the host population ear- 
ly in caterpillar development, and then the 
tachinid flies take their portion after that. 

The four species of dilleniaceous plants 
used by Enyo ocypete in this dry forest hab- 
itat are also fed on by caterpillars of the 
small sphingids /l/t'i/ro/z iphis, Llnzela japix, 
Unzela pronoe, and Pachygonidia drucei. 
However, of a total of 463 caterpillars of 
these species reared to date, none were 
parasitized by Euplectrus. These plants are 
also fed on by five species of Noctuidae, 
and no Euplectrus have been found on 
these caterpillars. 

In summary, Euplectrus floryae is un- 
ambiguously a specialist on Enyo ocypiete 
in this dry forest, but it uses at least four 
other species of very abundant sphingid 
caterpillars, none of which feed on the 
Enyo ocypete host plants, at a low frequen- 
cy. It is "ignoring" in some sense at least 
50 other species of sphingid caterpillars 
and a thousand or more species of other 
caterpillars whose size would not pre- 
clude sucessful E. floryae development in 
this dry forest habitat. Like all other Eu- 
plectrus reared in the ACG, it has never 
been found in rainforest or cloud forest, 
either with traps or by rearing. 



Volume 10, Number 2, 2001 



191 



Eiiplcctriis irciiene Schauff. — This species 
has been reared only once out of 29 rear- 
ings, from a rare and highly seasonal 
small green noctuid caterpillar, Motya ah- 
seuznlis L. This caterpillar feeds on the 
leaves of Couocnrpnis crccta (Combretaceae) 
during the second month of the rainy sea- 
son in the ACG dry forest at the edge of 
the coastal mangrove forest. Euplcciriis ir- 
eneae uses its last instar caterpillar host the 
same way as does Euplcctnis iiiiirinc. This 
host caterpillar, and the other caterpillars 
on the edges of the ACG mangrove 
swamps, have not yet been censused suf- 
ficiently to be able to say anything about 
the relative abundance or specificity of Eii- 
plectnis irenene. 

Euplectnis ivonae Schauff. — This species 
has been reared only twice out of 174 rear- 
ings of an extremely abundant and highly 
seasonal noctuid Eiiscirrlioptcnis poci/i. 
This caterpillar feeds on Pisoiiin iiincrniitlio- 
carpn Donn.-Sm. (Nyctaginaceae) only 
during the first week before and after the 
rainy season begins. This small eulophid 
wasp may either occur at an extraordi- 
narily low density or it normally uses 
some other species of caterpillar that has 
not yet been censused in the caterpillar in- 
ventory and feeds on some other host 
plant Euscirrhopterus pocyi Groteand two 
very low density leaf-rolling pyralids, Psa- 
ra hesperialis and Psara pnminidcs, are the 
only species of caterpillars feeding on Pi- 
soiiin iiincrniithocnrpm Donn.-Sm. in the 
ACG dry forest. 

Euplcctnis josci Schauff. — This species 
has a relationship to its sole host, Pacctcs 
hinodcs (Noctuidae), that is essentially 
identical to that of Eiip^lcctnis marine to its 
primary host. 

Eiiplcctrus magdac Schauff. — This species 
occurs at very low density on Dasylophia 
spp. (Notodontidae) in dry forest (Table 
1). It shares this host genus with Eiiplcctrus 
mariac, which is much more abundant, 
however on other caterpillars. 

Euplcctrus mariac Schauff. — This species 
appears to be a monophagous specialist 



on penultimate instar larvae of Coucaun 
muudissirua, a medium-small noctuid that 
feeds on three species of Malpighiaceae 
{Byrsonima crassifolia (L.) Kunth in HBK, 
Hiraea rcclinatn Jacq., Hetcroptcrys laurifolia 
(L.) A. Juss. in the ACG dry forest and on 
penultimate instar Elymiotis attcnuata (No- 
todontidae) also feeding on Malpighiaceae 
(Table 1). The few records of parasitizing 
last instar larvae may be correct, or may 
have been penultimate instar larvae. Its bi- 
ology of host use is essentially identical to 
that of Euplcctrus floryae, except that Eu- 
plcctrus mariac has been found only in the 
May-June first two months of the rainy 
season {Coiicaua muudissima also breeds al- 
most entirely during the first three months 
of the rainy season). There is no hint of 
pupal dormancy by Euplcctrus mariac, so 
we assume that it passes the reniainder of 
the year as a reproductively dormant 
adult or migrates to a wetter area to the 
east to have further generations. 

There are 37 records of Euplcctrus mariac 
from about 150 suitable-sized caterpillars 
of Cojicaua muudissima captured to date. 
No other noctuid caterpillar resembling, 
or taxonomically similar to, Coucaua muu- 
dissima is attacked by any species of Eu- 
plcctrus in this forest. This small Euplcctrus 
(5-10 larvae per caterpillar) could conceiv- 
ably parasitize any one of at least a thou- 
sand species of caterpillars living in the 
habitat of Coucaua unuuiissima. 

Euplcctrus mariac shares Concauua muu- 
dissima with 4 species of microgastrine 
braconid wasps and four species of tachi- 
nid flies. All together, these parasitoids 
take about 35% of the Coucaua muudissima 
caterpillars (which is exceptionally high 
for this inventory), and Euplcctrus mariae 
is responsible for about half of this cater- 
pillar mortality. 

Euplcctrus waltcri Schauff. — This is a low 
frequency parasitoid of penultimate (usu- 
ally) and ultimate (rarely) larvae of vari- 
ous species of Mauduca (Sphingidae) (Ta- 
ble 1). There has been one or more rearing 
of Euplcctrus zualtcri from all five of the 



192 



Journal o? Hymenoptera Research 



common Maiidiica species in the ACG dry 
forest. The absence of Eiiplectrus ivnlteri 
from the two rare species (Mnuducn sexta, 
Mnndiicn hanuibnl) is probably due to the 
very low numbers of these caterpillars 
found to date. The single record of Eiiplec- 
trus walteri from 381 caterpillars captured 
of Perigonia ilus undoubtedly represents 
an "abnormal" host record. These records 
indicate that either Eiiplectrus zvalteri is ig- 
noring the more than 50 other species of 
sphingid caterpillars in this habitat (more 
than 15,000 rearing records), or is unable 
to develop in them (though the single Per- 
igonia ilus record implies that the latter is 
unlikely). The larvae and cocoons of Eii- 
plectrus walteri are so distinctive that it can 
be stated with certainty that they are not 
using the caterpillars of any other species 
in the ACG dry forest (based on a sample 
of more than 50,000 caterpillars large 
enough to potentially host at least a small 
group of Eiiplectrus walteri larvae) 

A single Eiiplectrus walteri (Fig 4) lays 
up to several hundred eggs on the back of 
a (usually) penultimate instar Maudiica, 
and the larvae develop into a large patch 
of fat elongate green larvae tightly packed 
into one feeding area (Fig 6). hi all cases, 
all the larvae have been bunched together 
in one place, as if all the eggs were laid 
there by a single female. In the one case 
of oviposition observed, there were two 
females on the caterpillar (Fig 4) but only 
one was observed to oviposit. As with 
other Eiiplectrus larvae, they are strongly 
attached to the caterpillar cuticle by their 
mouthparts. Before the end of the cater- 
pillar's penultimate instar they have de- 
veloped to full size, released their hold, 
and spread into a ruff around the mori- 
bund caterpillar as they spin their cocoons 
(Fig. 7). The cocoons are stuck to one an- 
other and constitute a tight brown thick 
ruff around the caterpillar, which is dead 
but still clinging to the host plant leaf by 
the proleg crochets. When rearing this 
species, it is important not to disturb the 
caterpillar or wasp larva at the time that 



they release their hold on the caterpillar 
and move to their spinning site, as they 
easily fall off the caterpillar, become dis- 
oriented, and die. The wasps eclose over 
a 1-2 day period 10-14 days after spin- 
ning. There has been no suggestion of pu- 
pal dormancy during any of the 16 rear- 
ings. 

Eiiplectrus walteri has been found para- 
sitizing Maudiica from June through No- 
vember. Maudiica caterpillars are absent 
from the ACG dry forest habitat for the 
other months of the year and are extreme- 
ly rare after July. Either they pass the end 
of the rainy season and the dry season as 
reproductively dormant adults, or they 
migrate to the wetter portions of the ACG 
and points further to the east in the Carib- 
bean lowland rainforest. However, we fa- 
vor the former hypothesis since there are 
to date neither rearings nor Malaise trap 
records of Eiiplectrus walteri from any wet 
portion of Costa Rica. 

Eiiplectrus zvalteri occurs at low frequen- 
cy (32 cases out of more than 230 penul- 
timate instar records), and shares its Mau- 
diica hosts with a microgastrine braconid 
{Microplitis sp.), three species of ichneu- 
monidae (Janzen and Gauld 1997), and six 
species of Tachinidae in the ACG dry for- 
est. It is important to recall that if the cat- 
erpillar is collected prior to the penulti- 
niate instar, it cannot have E. walteri in it. 
In the three cases where Eiiplectrus walteri 
appeared to have attacked a last instar cat- 
erpillar, it could either be an exceptional 
record or the caterpillar was incorrectly 
determined to be a last instar. 

In addition to being the largest of the 
Eiiplectrus species reared in this study, Eii- 
plectrus walteri produces 10-20 times the 
number of wasps per caterpillar attacked 
as do the other species, and attacks the 
largest caterpillars known to be attacked 
by any species of Eiiplectrus. Eiiplectrus 
walteri also uses 3-6 days longer in the pu- 
pal stage than do the other smaller species 
of Eiiplectrus. 

Eiiplectrus xioniarae Schauff. — This spe- 



Volume 10, Number 2, 2001 



193 



cies is a specialist on early instar larvae of 
Hemiceras and Rosenin (Notodontidae) 
feeding on Iiign (Fabaceae). It has not been 
found on any of the tens of species of oth- 
er caterpillars feeding on Inga in the same 
dry forest. 

TAXONOMY 

Euplectrus anae Schauff, new species 

(Figs. 34, 40-42) 

Diagnosis. — Face below and between to- 
ruli yellow, not extending to eye or past 
gena (Fig. 13); legs yellow; one pair of se- 
tae ss2 between lateral ocelli (as in Fig. 70); 
all setae Sl-6 present (see Fig. 23); longi- 
tudinal carina on niesoscutum nearly 
complete, midlobe without small setae; 
scutellum finely reticulate; propodeum 
laterad of median carina nearly smooth; 
petiole wider than long. 

This species is similar to E. iiingdae 
which also has all the major setae on the 
face present. It can be differentiated by the 
petiole being longer than wide whereas 
related species all have the petiole as wide 
as long or wider than long. 

Description. — Female. Body length 
2.25mm. Color: body mostly black except 
the following: face below and between to- 
ruli yellow, not extending to eye or past 
gena; antenna with scape white to light 
yellow, flagellum yellow; mandibles 
white; legs yellow; dorsal metasoma with 
large central yellow area extending from 
just behind petiole posteriorly for about 
2/3 length, becoming darker in posterior 
1/4, laterally dark brown; ventral meta- 
soma yellow. Head. Dorsally with one pair 
of minor seta ss2 between posterior ocelli 
(as in Fig. 70) inserted near occipital cari- 
na; seta S5 reduced, setal row sr2 present 
as 2-3 irregular rows of 10-15 setae reach- 
ing the bottom of the eye; occipital carina 
present medially; width of eye: width of 
face 12:35; posterior margin of eye sepa- 
rated from margin of head ventrally. Ratio 
of MS:EH 15:29; lateral ocellus more than 
1 diameter from eye (OD:OOD 12:10). 



Face below eyes rounded, not abruptly 
narrowing. Vertex under anterior ocellus 
reticulate to alutaceous. Toruli separated 
by about 2-2. 5x their own diameters. Ma- 
lar suture absent. Area under eye irregu- 
larly reticulate to alutaceous. Scape 4X as 
long as wide. Ratio of funicular segments 
12:11:11:11:16, width 6 at Fl to 7 at club, 
each flagellar segment with scattered 
semierect brown setae, not arranged as 
whorls basally. Mesosonia. Pronotum an- 
terior to transverse carina with scattered 
setae, finely rugosely reticulate, posterior 
to carina more openly reticulate and 
shiny. Mesoscutum (Fig. 42) rugosely re- 
ticulate in anterior 1/2, becoming more 
open, smooth, and shiny posteriorly. Mid- 
lobe with median carina fading in anterior 
1/4 otherwise complete, slightly sunken 
posteriorly, posterior setae even with sur- 
face, with no sn^iall setae antero-laterally. 
Dorsal axillar/scutellar margin with 
broad, nearly straight deep furrow with 
flat bottom. Axillae shiny, openly reticu- 
late, becoming smooth at posterior mar- 
gin. Scutellum finely reticulate and shiny 
medially to more finely and striate retic- 
ulate laterally, pointed at anterior margin 
with axillae. Metanotum bordered anteri- 
orly and medially by small alveoli, medi- 
ally expanded into a triangular flange 
(Fig. 40) with submedian carinae below. 
Propodeum laterad of median carina 
sunken and irregularly carinate alveolate, 
becoming nearly smooth laterally to the 
step-like plica, median carina with anteri- 
or cup-like flange rounded and deeply in- 
vaginated. Area around spiracle finely re- 
ticulate, lateral edge of spiracle raised 
above surface, with antero-lateral flange 
large and well defined, with 6-8 setae lat- 
erad and below spiracle. Posterior margin 
of propodeum with irregular alveolae and 
carinae. Petiole in dorsal \'iew wider than 
long (15:10) and rugose dorsally becoming 
smooth at posterior margin. Metasoma. 
Ovate, about 1.5X as long as wide, with 
continuous brown margin laterally. Legs. 
Ratio of hind tibial spur l:spur 2:tarsus 1: 



194 



Journal of Hymenoptera Research 



T2:T3:T4. 35:23:22:10:8:20. Forewing. Hya- 
line, about 2.1 X as long as wide. Costal 
cell with 2 irregular rows of setae ventral- 
ly. Venation yellow, ratio of postmarginal: 
stigmal 35:20. 

Mfl/e.-Similar to female except: body 
length 1.8 mm; face below toruli lighter 
yellow, almost white; legs yellow or 
white; dorsal metasoma with large central 
white spot, dark brown posteriorly; anten- 
na with scape white slightly swollen on 
ventral surface, with sensory area slightly 
darker and with several irregular rows of 
sensillae extending for about 3/4 length 
(Figs. 34, 41); funicle ratios 11:10:10:10:15, 
width about 6 anteriorly to 7 posteriorly, 
with numerous semi-erect brown setae on 
each flagellomere. 

Hosts. — Sphncelodes vubicria (Geometri- 
dae) 

Distribution. — Known only from the 
ACG. 

Types. — Holotype female on point: Cos- 
ta Rica, Guanacaste Prov., Area de Con- 
servacion Guanacaste, Lanibert N309450 
E355300, 10 m., V. 11, 1992, 92-SRNP-747, 
D.H. Janzen & W. Hallwachs. ex. Sphnce- 
lodes vuineraria (Geometridae), (deposited 
in INBIO). Paratypes: 3 females and 2 
males with the same data as holotype (de- 
posited in USNM). 

Etymology. — This species is named in 
honor of Ana Leticia Martinez Eras in spe- 
cial recognition of her dedicated attention 
to the Accounting Office for the Area de 
Conservacion, Guanacaste. 

Euplectrns carloivae Schauff, new 
species 

(Figs. 25, 43) 

Diagnosis. — Face under toruli black; all 
legs, including hind coxae, yellow; first fu- 
nicle about sanie length as antennal club 
(Fig. 25); posterior margin of scutellum ex- 
tended over anterior margin of metano- 
tum laterally, anterior edge of metanotum 
medially expanded outward and divided 
into two areolae; hind basitarsus about 
equal in length to second tarsomere. Eii- 



plectrus carlowne is unusual in ha\'ing the 
hind basitarsus nearly equal in length to 
the second tarsus, while in all other spe- 
cies exaniined the first tarsomere is much 
longer than the second. It is also some- 
what unusual in having a dark face and 
all yellow legs (although this is shared 
with £. editliae). Other species treated here 
with the face dark brown or black have at 
least the hind coxa darkened {~onl-oerdei, za- 
nwrni, xioniarnc). The lateral expansion of 
the scutellum over the anterior edge of the 
metanotum is also distinctive. 

Description. — Female. Body length 2.2 
mm. Color: body mostly black except the 
following: antenna with scape yellow to 
brown, flagellum light brown becoming 
darker brown apically; mandibles light 
brown; enlarged setae on vertex dark 
brown to black; legs yellow; dorsal meta- 
soma dark brown to black behind petiole 
with yellow inverted T-shaped spot me- 
dially, posterior half dark brown, ventral 
metasoma dark brown behind petiole, 
then yellow" up to about midpoint, then 
dark brown. Head. Dorsally with 2 minor 
seta ss2 between posterior ocelli, all setae 
Sl-6 present, setal row sr2 present as 2 ir- 
regular rows of about 10 setae reaching to 
bottom of eye; occipital carina weak me- 
dially; width of eye: width of face (30:13), 
posterior margin of eye not nearly contig- 
uous with posterior margin of head over 
most of length. Ratio of MS:EH 17:29; lat- 
eral ocellus more than 1 diameter from 
eye (OD:OOD 7:8). Face below eyes 
rounded, not abruptly narrowing. Vertex 
under anterior ocellus reticulate. Toruli 
separated by about 2x their own diame- 
ters. Malar suture absent. Area under eye 
lightly reticulate. Scape 5.5 X as long as 
wide. Ratio of funicular segments 18:17:15: 
13:18, width 5 at Fl to 6 at club, flagellar 
segments with small whorls of brown se- 
tae basally. Mesosoma. Pronotum anterior 
to transverse carina with scattered setae, 
finely rugosely reticulate, posterior to ca- 
rina reticulate and shiny. Mesoscutum 
(Fig. 43) reticulate, becoming more open. 



Volume 10, Nl'mbfr 2, 2001 



195 



W 



% 






Figs. 10-17. Eiiplt'ctrus. 10-11, Hindlegs. 10, E.florytw. 1 1, £. xiowarae. 12-16, Head, frontal view. 12, E.flon/ae. 
13, £. -cvaltcri. 14, £. //r;uv?('. 15, E. arias. 16, £. /j'onrtc. 17, Male scape, £. valwrdci. 



196 



Journal of Hymenoptera Research 



smooth, and shiny posteriorly. Midlobe 
with median carina fading in anterior 1/4, 
not noticeably sunken posteriorly, with no 
small setae antero-laterally, posterior setae 
even with surface. Dorsal axillar/scutellar 
margin with broad, distinctly curved deep 
furrow with flat bottom. Axillae openly 
reticulate. Scutellum reticulate to aluta- 
ceous, smooth along posterior margin, 
posterior n^iargin extended over anterior 
margin of metanotum laterally. Metano- 
tum laterally covered by scutellum, me- 
dially with two distinct protruding alve- 
oli, below shiny and smooth to lightly re- 
ticulate. Propodeum laterad of median ca- 
rina shiny and smooth to the step-like 
plica with occasional faint hints of reticu- 
lation, median carina distinctly raised 
above surface, with large, rounded ante- 
rior cuplike flange. Area around spiracle 
reticulate, spiracle slightly raised and ev^en 
with surface, with antero-lateral flange 
present, with 7 setae laterad and below 
spiracle. Petiole in dorsal view as long as 
wide (12:11), rugose dorsally. Mctasonin. 
Ovate, about 2x as long as wide. Legs. Ra- 
tio of hind tibial spur l:spur 2:tarsus 1:T2: 
T3:T4. 38:22:17:17:12:18. Forewing. Hya- 
line, about 2.4 X as long as wide. Costal 
cell with 1 irregular row of setae ventrally. 
Venation yellowish, ratio of postmarginal: 
stigmal 31:16. 

Male. — Unknown. 

Hosts. — Unknown. 

Distribution. — Known only from Puntar- 
enas and Alajeula. 

Types. — Holotype female on point (an- 
tenna and wing slide-mounted) with data: 
"Costa Rica, Puntarenas, R. F. Golfo Dul- 
ce, 24 Km. W. Piedras Blancas, 200 m., IV- 
V 1992. P. Hanson." (deposited in INBIO). 
Paratypes: 2 females with same data as 
holotype except II-III. 1989, 2 females XII. 
1989-III. 1990, and 2 females at 100m, III- 
V. 1989; 1 female Costa Rica: Alajuela, 
5km W. San Ramon, 1200m, I. 1997, O. 
Castro & P. Hanson (deposited in USNM, 
BMNH). 

Eti/iiioh\^if. — This species, collected only 



from Malaise trapping, is named for Ms. 
Tami Carlow of the Systematic Entoniol- 
ogy Lab, USDA who was responsible for 
most of the scanning electron micrographs 
used in this paper and also assisted with 
specimen mounting, labelling and a vari- 
ety of other tasks vital to the completion 
of this work. 

Etiplectrus comstockii Howard 

(Figs. 27, 44) 

Eiiplcctnis coiustockii Howard 1880:158. 

Diagnosis. — Legs yellow; face under to- 
ruli yellow; first funicle 2x as long as 
wide; seta 5 present (see Fig. 23), but re- 
duced; with one pair of small setae (ss2) 
between posterior ocelli (as in Fig. 66); Fl 
2x as long as wide; median longitudinal 
mesoscutal carina nearly complete; scutel- 
lum lightly reticulate and shiny medially, 
becoming striate laterally; anterior metan- 
otum with large, obvious line of alveoli 
(Fig. 44), central transverse band narrow, 
smooth; propodeum adjacent to median 
carina lightly reticulate, shiny. Petiole 
wider than long. Metasoma mostly yel- 
low, becoming brown posteriorly. Male 
scape slightly (Fig. 27) expanded near 
apex, with 2 irregular short rows of sen- 
sillae. 

£. comstockii is most siniilar to solitiiriiis 
but can be separated by the petiole which 
is longer than wide in solitarius and wider 
than long in coiustockii and the antennae 
which has Fl 2X as long as wide in coiu- 
stockii and 3x as long as wide in solitarius. 

Distribution. — Widespread in United 
States, Central and South America. 

Hosts. — The following records are pri- 
marily drawn from the literature (Burks 
1979, Noyes 1998). In a few instances they 
have been verified from specinien label 
data in collections (USNM, CNC). It is our 
opinion, based on data from the species 
reared in this study, that at least some of 
these records probably represent misiden- 
tifications of either the parasite or the host. 
Alypia octoiuaculata; Anoiuis iUita; Autogra- 



Volume 10, Number 2, 2001 

Postmarginal Vein 



197 





Stiemral Vein . 



" Measure froni 
A-BorC-D- 






Figs. 18-22. EupUxtnis. 18, Forewing \cnatiun. 1^), £. xiiviiarac, male scape,. 20-21, E. valverdei, male scape. 
22, £. zainorai, head and antennae. 



pha sp.; Caradrina sp.; Hadciia luieago; Hcl- 
icoverpa armigera; Helicoverpm zca; Hcliothis 
sp.; Heliothis viresceus; Hi/pcim scnbm; Lcit- 
cnuin latiiiscula; Ncognica siDiia; Pliisin sp; 
Pscudopliisin indudcns; Sclciiisn siieroidcs; 
Spodoptcrci frugipcrda; Spodoptern oniitliogal- 
li; Trichoplusin ni (Noctuidae); FaiwndcUn 
fiiiictnrin (Geometridae); Rothschildn nnviin 
(Saturniidae). 

Typjcs. — Howard described this species 
from "two male specimens". He indicated 
that the original specimens had been col- 
lected by Comstock and that "upon look- 
ing them up, I found that two adults had 



issued". In the U.S. National Museum col- 
lection, there is a series containing both 
males and females, on cards and points 
and all bearing the USNM type no. 2653. 
The type catalog entry for this species lists 
both male and female specimens collected 
in 1878 and 1880 by W.H. Patton and S. 
A. Schwarz (not Comstock) and the num- 
ber of specimens listed is "many". How- 
ard made no mention of localities in his 
original description, but all the USNM 
specimens with locality labels are from 
Selma, Alabama, collected in 1880, except 
one collected in 1878 from Florida. Ac- 



198 



Journal of Hymenoptera Research 



cording to the description, the types were 
reared in 1879 during fieldwork conduct- 
ed by Comstock. None of the points or 
cards contains only 2 males. Given these 
facts, we believe it likely that none of the 
specimens labelled as types in the USNM 
collection are in fact the specimens that 
Howard used for his original description, 
but rather the specimens currently la- 
belled as types are specimens used by him 
for a subsequent redescription (Howard 
1885). Since the original type specimens 
are lost, we are erecting a neotype (present 
designation). This specimen is one of the 
series labelled as types. It is a female on a 
card with 3 other specimens and has been 
marked with an "N" in black ink. The la- 
bel data is: "Selma, Oct., 81. Patton. Type 
no. 2653, U.S.N.M." 

Euplectrtis edithae Schauff, new species 

(Fig. 39) 

Diagnosis. — Face black below toruli (as 
in Fig. 15), legs yellow; one pair of setae 
ss2 between lateral ocelli; flagellum 
brown, funiculars about 2x as long as 
wide; malar space nearly equal to eye 
height; petiole as wide as long; postmar- 
ginal less than 1.5 X stigmal. This combi- 
nation of a black face below the toruli and 
yellow hind coxae is found only in this 
species and in E. cnrloivne. In addition, E. 
carlowae has the second tarsomere of the 
hind leg nearly equal in length to the first 
(second tarsomere much shorter than first 
in cdithac) and the lateral scutellum is ex- 
panded and overlaps the metanotum (lat- 
eral scutellum not overlapping metano- 
tum in edithae). 

Description. — Female. Body length 2.2- 
2.3 mm. Color: body mostly black except 
the following: antenna with scape yellow 
to brown, flagellum brown; mandibles 
yellow; enlarged setae on vertex yellow to 
dark brown; legs yellow; dorsal metasoma 
dark yellow behind petiole and ventrally, 
dark brown laterally. Head. Dorsally with 
2 minor seta ss2 between posterior ocelli, 
seta S5 reduced, setal row sr2 present as 2 



irregular rows of about 8 setae reaching to 
bottom of eye; occipital carina reduced; 
width of eye: width of face (40:13), poste- 
rior margin of eye not nearly contiguous 
with posterior margin of head over niost 
of length. Ratio of MS:EH 19:22; lateral 
ocellus more than 1 diameter from eye 
(OD:OOD 5:10). Face below eyes abruptly 
narrowing. Vertex under anterior ocellus 
reticulate. Toruli separated by about 2x 
their own diameters. Malar suture absent. 
Area under eye lightly reticulate. Scape 
6X as long as wide. Ratio of funicular seg- 
ments 11:11:11:11:16, width 5 at Fl to 6 at 
club, flagellar segments with small whorls 
of brown setae basally. Mesosoina. Prono- 
tum anterior to transverse carina with 
scattered setae, finely rugosely reticulate, 
posterior to carina reticulate and shiny. 
Mesoscutum reticulate, becoming more 
open, smooth, and shiny posteriorly. Mid- 
lobe with median carina complete, not no- 
ticeably sunken posteriorly, with no small 
setae antero-laterally, posterior setae even 
with surface. Dorsal axillar/scutellar mar- 
gin with relatively narrow, parallel sided 
furrow making anterior margin of scutel- 
lum distinctly V-shaped. Axillae openly 
reticulate. Scutellum reticulate to aluta- 
ceous, smooth along posterior margin, 
posterior margin not extended over ante- 
rior margin of metanotum laterally. Me- 
tanotum with a line of alveoli anteriorly. 
Propodeum laterad of median carina 
lightly reticulate, shiny and smooth to the 
step-like plica, median carina distinctly 
raised above surface, with large anterior 
cup-like flange which is somewhat trun- 
cated posteriorly. Area around spiracle re- 
ticulate, spiracle slightly raised and even 
with surface, with antero-lateral flange 
present, with 10-12 setae laterad and be- 
low spiracle. Petiole in dorsal view as long 
as wide (10:10), rugose dorsally. Metasoma. 
Ovate, about 1.5 X as long as wide. Legs. 
Ratio of hind tibial spur l:spur 2:tarsus 1: 
T2:T3:T4. 37:25:25:13:10:13. Forewing. Hy- 
aline, about 2.5 X as long as wide. Costal 
cell with 2 irregular rows of setae ventral- 



Volume 10, Number 1, 2001 

ssl S2 ss2 



199 




24 



HW 



Figs. 23-26. Euplectnis morphology. 23, Head in dorsal view. 24, Head in frontal view. 25, Female antenna 
of £. carlowae. 26, Female antenna of E. hansom. EW = eye width. EH = Eye height. FW = Face width. FH = 
Face height. HW = Head width, MS = Malar space. OOD = Ocell-ocular distance. OOL = Ocell-ocular length. 
POL = Posterior ocellar length. Sl-6 = Major setae 1 to 6. sr = setal row. ss = small setae. OD = ocellar 
diameter. 



ly. Venation yellowish, ratio of postmar- 
ginal: stigmal 32:25. 

Male. — Similar to female except: scape 
yellow to white and slightly swollen api- 
cally (Fig. 39) with two irregular rows of 
sensillae running about 2/3 length; legs 
generally yellow; metasoma darker, near- 
ly black in posterior half both dorsally and 
ventrally; funicle ratio 11:11:11:10:15, 
width 4-6 with no noticeable brown setae. 

Since the male of this species was not 
reared with the associated females, I can- 
not be absolutely positive of the relation- 



ship. However, all the specimens were col- 
lected at the same locality at the same time 
and although other species were also pre- 
sent in those collections, this male matches 
the females of this species much more 
closely, and we are sufficiently confident 
of the association to assign it to this spe- 
cies. 

Has ts. — U n kno vv n . 

Distribution. — Costa Ri ca . 

Types. — Holotype female with data: 
"Costa Rica, San Jose, Zurqui de Moraxia, 
1600m, IV. 1995, P. Hanson", (deposited in 



200 



Journal of Hymenoptera Research 



INBIO). Paratypes: 10 females and 1 male 
with same data except 2 collected in Jan- 
uary 1996 (deposited in USNM and 
BMNH). 

Eti/iJioIogi/. — This species is named in 
honor of Edith Lopez Lara in special rec- 
ognition of her dedicated attention to the 
Research Center and Dormitories in Sector 
Santa Rosa of the Area de Conservacion 
Guanacaste. 

Etiplectnis floryae Schauff, new species 

(Figs. 10, 12, 28, 45-48) 

Diagnosis. — Face below and between to- 
ruli yellow, extending up side of eye to 
midpoint and around face to gena and 
mouth (Fig. 12); two pairs of small setae 
(ss2) between lateral ocelli (Fig. 66); me- 
soscutal midlobe with 1-2 small setae an- 
teriorly (Fig. 48), median carina complete; 
scutellum heavily reticulate to alutaceous; 
petiole in dorsal view as long as wide and 
rugose dorsally with irregular longitudi- 
nal carina. Male antennal scape white, 
slightly swollen, with narrowly ovate sen- 
sory area containing 2-3 irregular rows of 
sensillae extending about 3/4 length (Fig. 
28), Fl slightly shorter than club. 

The coloration of the face with yellow 
running up the side of the eyes makes this 
species quite distinctive among Costa Ri- 
can species. In addition, two pairs of small 
setae between the lateral ocelli and small 
setae on the anteric^r mesoscutal midlobe 
distinguish it from similar species treated 
here. This species is similar to Eiiplcctrns 
ijinculivciitris Westwood which is wide- 
spread in Canada and the U.S. and which 
may occur in Central America. E. luaculi- 
vcutris has the face yellow with the yellow 
extending up the side of the eyes. Flow- 
ever, E. Jiiaciilivciitris has the median cari- 
na on the scutum nearly absent and the 
antenna of the male has the scape more 
enlarged and with 3-6 rows of sensillae, 
and the funicles are elongate and each is 
covered by elongate setae. 

Description. — Female. Body length 2.1- 
2.3 mm. Color: body mostly black except 



the following: face below and between to- 
ruli yellow, extending laterad of toruli 
over gena, down to mouth, and up edge 
of eye to near midpoint (Fig. 12); antenna 
with scape white to light yellow, flagellum 
yellow or light brown; mandibles yellow 
to white; legs light yellow to white; dorsal 
metasoma with large central yellow to 
white area extending from just behind pet- 
iole posteriorly over entire length of dor- 
sum, becoming slightly darker posteriorly, 
lateral brown margin reduced to two 
brown spots separated medially by yellow 
and ending well before posterior margin, 
ventral metasoma yellow to white. Hcnd. 
Dorsally with two pair of minor setae ss2 
between posterior ocelli (as in Fig. 46), in- 
serted adjacent to occipital carina, setae 
Sl-6 present (as in Fig. 23), setal row sr2 
present as 1-2 irregular rows of 12-16 se- 
tae, reaching the bottom of the eye. Occip- 
ital carina strongly present over entire 
length of occiput. Width of eye: width of 
face 13:35, posterior margin of eye nearly 
contiguous with posteric^r margin of head 
of most of length. Ratio of MS:EH 17:31, 
lateral ocellus more than 1 diameter from 
eye (OD:OOD 6:8). Face below eyes 
rounded, not abruptly narrowing. Vertex 
below anterior ocellus reticulate to aluta- 
ceous. Toruli separated by about 2-2.2 X 
their own diameters. Malar suture absent. 
Area under eye lightly reticulate to 
smooth (difficult to assess because of yel- 
low coloration). Scape 5x as long as wide. 
Ratio of funicular segments 14:12:12:13:17, 
width 5 at Fl to 6 at club, flagellar seg- 
ments with small indistinct whorls of 
brown setae basally. Mesosouw. Pronotum 
anterior to transverse carina with scat- 
tered setae, finely rugosely reticulate, pos- 
terior to carina more openly reticulate and 
shiny. Mesoscutum (Fig. 48) reticulate, be- 
coming more smooth, and shiny posteri- 
orly. Midlobe with median carina well de- 
fined, fading only at extreme anterior mar- 
gin, otherwise con^plete, slightly sunken 
posteriorly, with one or two small setae 
antero-laterally, posterior setae even with 



Volume 10, Number 2, 2001 



201 



surface or slightly raised. Dorsal axillar/ 
scutellar margin with broad, curved deep 
furrow with flat bottom. Axillae shiny, 
openly reticulate. Scutellum shiny and 
lightly reticulate to alutaceous. Metano- 
tum bordered anteriorly by a narrow band 
of small alveoli, medially shiny and light- 
ly reticulate. Propodeum laterad of medi- 
an carina shiny and openly reticulate to 
the step-like plica, median carina with an- 
terior cup-like flange rounded and invag- 
inated. Area around spiracle finely retic- 
ulate or granulate, spiracle slightly raised 
above surface (Fig. 47), opening parallel to 
the surface of the propodeuni, with an- 
tero-lateral flange large and obvious, with 
10-12 setae laterad and below spiracle. 
Petiole in dorsal view as long as wide (10: 
10), rugose dorsally with irregular longi- 
tudinal carina. Metasoina. Ovate, about 
1.5-2X as long as wide. Legs. Ratio of 
hind tibial spur l:spur 2:tarsus 1:T2:T3:T4. 
35:25:23:13:10:13. Forewing. Hyaline, 
about 2.5 X as long as wide. Costal cell 
with 2-3 irregular rows of setae ventrally. 
Venation yellow to white, ratio of post- 
marginal: stigmal (28:18). 

Male. — Similar to female except: body 
length 1.8-2mm; face white, legs white to 
light yellow, metasoma with central white 
spot only extending about 1/2 to 2/3 of 
length, dark brown posteriorly, laterally 
dark brown, sometimes interrupted me- 
dially; antenna with scape white (Fig. 45), 
slightly swollen, with narrowly ovate sen- 
sory area containing 2-3 irregular rows of 
sensillae extending about 3/4 length (Fig. 
28); funicle ratios 13:12:14:12:15, width 5, 
without scattered or whorled semierect 
brown setae on each flagellomere. 

Hosts. — Ein/o ocypetc, Perigonia ilus, P. 
lusca, Aellopos fasus (all Sphingidae). 

Distribution. — Known only from the 
ACG. 

Eti/uiology. — This species is named in 
honor of Flory Granados Venegas in spe- 
cial recognition of her dedicated manage- 
ment of the main Administrative Office of 
the Area de Conservacion Guanacaste. 



fi/pes. — Holotype female: "Costa Rica, 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N320050 E365300, 
160m., in. 5, 1992, 92-SRNP-524, D. H. Jan- 
zen & W. Hallwachs. ex. Eni/o ocypete" 
(deposited in INBIO). Paratypes: 3 females 
and 1 male all with the same data as ho- 
lotype; other specimens from the ACG: 2 
males with Lambert N317200 E360850, 
290m., V. 3. 1994, 94-SRNP-918; 2 males 
with Lambert N325500 E360200, 270m., 
VI. 14. 1990, 90-SRNP-n2; 2 females and 
1 male with Lambert N313800 E359800, 
300m., Vlll. 4. 1993, 93-SRNP-4177, ex. 
Perigoiiia ilus; 1 female anci 1 male with 
Lambert N319000 E361150, 270m., X. 11. 
1992, 92-SRNP-5353; 1 female with Lam- 
bert N319500 E360650, 260m., VI. 8. 1992, 
92-SRNP-1115; 1 male and 1 female with 
Lambert N314500 E357850, 290m., Vll. 27. 
1990, 90-SRNP-1522 ; 1 female with Lam- 
bert N314500 E357850, 290m., VI. 8. 1992, 
92-SRNP-1084; 1 male with Lambert 
N319550 E360650, 290m., VI. 10. 1992, 92- 
SRNP-1112; 2 females with Lambert 
N314500 E357850, 290m., Vlll. 5. 1992, 92- 
SRNP-3929; 1 male with Lambert N319100 
E360900, 260m., V. 25. 1992, 92-SRNP-831; 
1 female with Lambert N314500 E357850, 
240m., Vll. 21. 1993, 93-SRNP-2502, ex. 
Perigonin lusca; 1 male with Lambert 
N315700 E354400, 300m., VI. 6. 1992, 92- 
SRNP-1009; 1 female with Lambert 
N315500 E360200, 300m., Vll. 23. 1991, 91- 
SRNP-1792, ex. Pcrigouia ilus; 1 male with 
Lambert N319000 E361150, 270m., II. 2. 
1992, 92-SRNP-293, ex. Aellopos fad us; 1 fe- 
male with Lambert N314800 E360500, 
300m., VII. 23. 1984, 84-SRNP-1479; 1 male 
and 1 female with Lambert N314650 
E361300, 270m., XI. 20. 1993, 93-SRNP- 
7708 ex. Perigonia ilus; 3 males with Lam- 
bert N313100 E359900, 250m., VIII. 24. 
1990, 90-SRNP-1946; 1 female with Lam- 
bert N319100 E360900, 260m., VII. 13. 
1992, 92-SRNP-2612; 1 female with Lam- 
bert N312300 E361050, 260m., VI. 12. 1992, 
92-SRNP-1073; 1 female with Lambert 
N314500 E357850, 290m., V. 26. 1991, 91- 



202 



Journal of Hymenoptera Research 




Figs. 27-39. Euplcctriis male scapes. 27, E. coiiistockii. 2S, E. flonjac. 29, £. waltcri. 30, £. maviac. 31, £. romuci. 
32, £. ivoiKw. 33, £. iiias^dnc. 34, £. «;;(;t'. 35, £. /dst'/. 36, £. xionuinu-. 37, £. om/s. 38, £. vnlvcrdci. 39, £. cdiljmc. 



Volume 10, Number 2, 2001 



203 



SRNP-328; 2 males with Lambert N319000 
E361150, 270m., V. 10. 1994, 94-SRNP-935; 
1 female with Lambert N313800 E359800, 
300m., VI. 19. 1988, 88-SRNP-229, ex. Pcr- 
igouia ilus; 1 female with Lambert N314800 
E360500, 300m., VIII. 12. 1991, 91-SRNP- 
2479, ex. Pcrigoiiin ilus; 2 males with Lam- 
bert N31450d E357850, 290m., V. 28. 1991, 
91-SRNP-248; 1 female with Lambert 
N313100 E359900, 250m., VII. 4. 1982, 82- 
SRNP-368, ex. Pcrigouin ilus; 1 female with 
Lambert N313800 E359800, 300m., VII. 28. 
1984, 84-SRNP-1501 ex. Cnutcthia spuria; 1 
male with Lambert N315500 E360200, 
300m., VII. 15. 1991, 91-SRNP-1512, ex. 
Perigonin ilus; 1 female with Lambert 
N314800 E360500, 300m., VIII. 13. 1992, 
92-SRNP-3473; 1 female with Lambert 
N317200 E360850, 290m., V. 28. 1991, 91- 
SRNP-280; 1 female with Lambert 
N317200 E360850, 290m., V. 27. 1991, 91- 
SRNP-278; 1 female with Lambert 
N317200 E360850, 290m., V. 27. 1991, 91- 
SRNP-277 (deposited in USNM, BMNH, 
and CNC). 

Eiiplectrtis ftiruins Walker 

(Figs. 49, 50) 

Eiiplectms furnius Walker 1843:48. 

Pachyscapha insularis Howard 1897:159. (Syn- 
onymy by Boucek 1977; see also Boucek in 
Desantis 1979). 

Dinguosis. — Apical flagellar segments 
generally darker than Fl (as in Fig. 22); 
legs yellow; face below toruli dark brown 
to black; with no small setae between pos- 
terior ocelli (Fig. 49); width of face nearly 
4x width of eye; posterior ocellus 2X its 
diameter from eye margin; scutellum 
nearly smooth, shiny, with some light re- 
ticulation; petiole wider than long; post- 
marginal vein barely longer than stigmal 
(25:20). Male antenna with scape dark 
brown, greatly enlarged (Fig. 50), and uni- 
fornily covered with sensillae, with funic- 
ular segments quadrate and F3 and 4 dark 
brown contrasting with Fl and 2 which 
are yellow. 



Euplectrus furnius is a distinctive species 
easily recognized by the dark, broad face, 
with ocelli 2 diameters removed from the 
eye, gradually darkening antennal flagel- 
lum in the female (even more marked in 
the males) and short postmarginal vein. 
The dark, greatly swollen scape of the 
male is also quite distinctive. Although 
other species of Euplectrus are known to 
have similar scapes, none of those treated 
in this study have a swollen scape that is 
also dark colored. 

Hosts. — The following records are pri- 
marily drawn from the literature (Burks 
1979, Noyes 1998). In a few instances they 
have been verified from specimen label 
data in collections (USNM, CNC). Hosts 
include Agrius ciiiguhitus (Sphingidae); 
Autichloris criphia (Arctiidae); Hndcun lutca- 
go; Helicoverpa zea; Lninproscmn iiuiicatn; 
Pseudoplusia indudens; Rnchisplusia mi; Spo- 
doptern eridauia; S. frugipcrda (Noctuidae). 

Distribution. — Known from Mexico 
south to Ecuador and Venezuela and west 
to the West Indies. 

Types. — The lectotype of E. furnius is in 
The Natural History Museum London (ex- 
amined). Lectotype and paralectotypes of 
£. insularis are in the USNM (examined). 

Euplectrus liansoni Schauff, new species 

(Fig. 26) 

Diagnosis- Face yellow below toruli (as 
in Fig. 13), legs yellow; one pair of setae 
ss2 between lateral ocelli; Fl^ all about 
3X as long as wide and nearly equal to 
club (Fig. 26), flagellum dark brown; post- 
marginal more than 2X stigmal (see Fig. 
18). 

This species is most easily distinguished 
by the elongate and dark brown funicle 
segments (Fl-4 about 3X as long as wide 
and Fl-3 nearly as long as club). In £. wal- 
tcri in which Fl is also almost as long as 
the club, the funiculars are only about 2x 
as long as wide and the segments are yel- 
low or light brown colored. In addition, 
the long postmarginal vein (more than 2X 
as long as the stigmal) is longer than in 



204 



Journal of Hymenoptera Research 



the other species where it is usually 2x as 
long as the stigmal or less. 

Description. — Female. Body length 2.2- 
2.6 mm. Color: body mostly black except 
the following: antenna with scape yellow 
to brown, flagellum dark brown; mandi- 
bles yellow; enlarged setae on vertex yel- 
low to dark brown; legs yellow; dorsal 
metasoma dark brown to black behind 
petiole with yellow spot medially, poste- 
rior half dark brown, ventral metasoma 
dark brown behind petiole, then yellow 
up to about midpoint, then dark brown. 
Head. Dorsally with 2 minor seta ss2 be- 
tween posterior ocelli (as in Fig. 23), all 
setae Sl-6 present, setal row sr2 present 
as 2 irregular rows of about 8 setae reach- 
ing to bottom of eye; occipital carina ob- 
vious medially; width of eye: width of face 
(36:13), posterior margin eye of not nearly 
contiguous with posterior margin of head 
over most of length. Ratio of MS:EH 18:30; 
lateral ocellus more than 1 diameter from 
eye (OD:OOD 6:9). Face below eyes 
rounded, not abruptly narrowing. Vertex 
under anterior ocellus reticulate. Toruli 
separated by about 2X their own diame- 
ters. Malar suture absent. Area under eye 
lightly reticulate. Scape 5x as long as 
wade. Ratio of funicular segments 19:19:19: 
17:21, width 6 at Fl to 7 at club, flagellar 
segments with small whorls of brown se- 
tae basally. Mesosomn. Pronotum anterior 
to transverse carina with scattered setae, 
finely rugosely reticulate, posterior to ca- 
rina reticulate and shiny. Mesoscutum re- 
ticulate, becoming more open, smooth, 
and shiny posteriorly. Midlobe with me- 
dian carina fading in anterior 1/4, not no- 
ticeably sunken posteriorly, with no small 
setae antero-laterally, posterior setae even 
with surface. Dorsal axillar/scutellar mar- 
gin with broad, distinctly curved deep fur- 
row with flat bottom. Axillae openly retic- 
ulate. Scutellum reticulate to alutaceous, 
smooth along posterior margin, posterior 
margin not extended over anterior margin 
of metanotum laterally. Metanotum with 
a very thin line of alveoli anteriorly. Pro- 



podeum laterad of median carina lightly 
reticulate, shiny and smooth to the step- 
like plica, median carina distinctly raised 
above surface, with large anterior cup-like 
flange which is somewhat truncated pos- 
teriorly. Area around spiracle reticulate, 
spiracle slightly raised and even with sur- 
face, with antero-lateral flange present, 
with 6-7 setae laterad and below spiracle. 
Petiole in dorsal view longer than wide 
(14:10), rugose dorsally. Mctasomn. Ovate, 
about 1.5X as long as wide. Legs. Ratio of 
hind tibial spur l:spur 2:tarsus 1:T2:T3:T4. 
40:25:22:17:12:16. Forewing. Hyaline, 
about 2.4 X as long as wide. Costal cell 
with 2 irregular rows of setae ventrally. 
Venation yellowish, ratio of postniarginal: 
stigmal (50:23). 

Mnle. — Unknown. 

Hosts. — Unknown. 

Distribution. — Known only from the 
type locality. 

Types. — Holotype female, deposited in 
USNM, with data: "Costa Rica, San Jose, 
Zurqui de Moravia, 1600m, IX.1996, P. 
Hanson". Paratype female with same data 
deposited in USNM. 

Etymology. — This species is named for 
the collector of the types, Paul Hanson, 
who also contributed many other interest- 
ing specimens to this study. 

Eiiplectriis ireneae Schauff, new species 

(Figs. 14, 51) 

Diagnosis. — Face below and between to- 
ruli yellowish brown, lighter than above 
toruli; legs light yellow to white; width of 
eye more than half width of face (Fig. 14), 
lateral ocellus less than 1 diameter from 
eye; with 1 pair of setae (ss2) between lat- 
eral ocelli; setal row sr2 present as 1-2 ir- 
regular rows of 5-12 setae usually not 
reaching the bottom of the eye. 

The structure of the head with the lat- 
eral ocellus less than 1 diameter from the 
eye and the eye itself more than half the 
width of the face (frontal view) make this 
species easily recognizable from all other 
species treated. 



Volume 10, Number 1, 2001 



205 




Figs. 40^7. Euplccirus scanning electron micrographs. 40-42, E. miac. 40, I'ropodeum. 41, Male antennae. 42, 
Mesosoma. 43, £. cnrloimc, Scutellum and propodeum. 44, £. coiii>tockii, mesosoma. 45-47, E. floryae. 45, Male 
scape. 46, Head, dorsal \iew. 47, Propodeum. 



206 



Journal of Hymenoptera Research 




54 I'A/ . 

Figs. 48-55. Eiiplectnis scanning electron micrograpiih. 4cS, L. jloninc, liorsal hicmisoiil 
Head, dorsal view. 50, Male scape. 51, E. iroicac, propodeum. 52-53, £. Ivoiuic. 52, 
Propodeuni. 54-55, £. josci. 54, Head, dorsal view. 55, Dorsal mesosoma. 



1. 44-50, /: 
Dorsal me 



. finiiliiti. 4^), 
sosoma. 53, 



Volume 10, Number 2, 2001 



207 



Description. — Female. Body length 2.3- 
2.6 mm. Color: body mostly black except 
the following: face below and between to- 
ruli yellowish brown; antenna with scape 
white tc^ light yellow, flagellum yellow; 
mandibles yellow; legs light yellow to 
white; dorsal metasoma with large central 
yellow area extending from just behind 
petiole posteriorly for about 1/2 length, 
and extending laterally around the side of 
the metasoma, posteriorly dark brown, 
ventral metasoma yellow. Head. Dorsally 
with one pair of minor seta ss2 between 
posterior ocelli (as in Fig. 70), inserted dis- 
tinctly above occipital carina, all setae Sl- 
6 present, setal row sr2 present as 1-2 ir- 
regular rows of 5-12 setae usually not 
reaching the bottom of the eye; occipital 
carina present niedially; width of eye: 
width of face 17:30, posterior margin eye 
of contiguous with posterior margin of 
head of most of length. Ratio of MS:EH 13: 
36; lateral ocellus less than 1 diameter 
from eye (OD:OOD 12:5). Face below eyes 
rounded, not abruptly narrowing. Vertex 
under anterior ocellus lightly reticulate. 
Toruli separated by about 2-2.5 X their 
own diameters. Malar suture absent. Area 
under eye lightly reticulate. Scape 5x as 
long as wide. Ratio of funicular segments 
16:15:16:16:20, width 6 at Fl to 8 at club. 
Mesosoma. Pronotum anterior to trans- 
verse carina with scattered setae, finely ru- 
gosely reticulate, posterior to carina more 
openly retculate and shiny. Mesoscutum 
reticulate in anterior 1/2, becoming more 
smooth, and shiny posteriorly with slight 
reticulation. Midlobe with median carina 
fading only at extreme anterior margin, 
otherwise complete, not sunken posteri- 
orly, with no small setae antero-laterally, 
posterior setae even with surface. Dorsal 
axillar/scutellar margin with broad, 
slightly curved, deep furrow with flat bot- 
tom. Axillae shiny, openly reticulate, be- 
coming nearly smooth at posterior mar- 
gin. Scutellum finely reticulate to aluta- 
ceous, shiny, slightly pointed at anterior 
margin with axillae. Metanotum not bor- 



dered anteriorly and medially by small al- 
veoli, medially flat and shiny and very 
lightly reticulate without median carina 
below. Propodeum laterad of median ca- 
rina nearly smooth (lightly reticulate) lat- 
erally to the step-like plica, median carina 
with anterior cup-like flange rounded and 
invaginated. Area around spiracle flnely 
reticulate, lateral edge of spiracle raised 
above surface (Fig. 51), with antero-lateral 
flange large and well defined, with 7-8 se- 
tae laterad and below spiracle. Posterior 
margin of propodeum without irregular 
alveolae and carinae. Petiole in dorsal 
view slightly wider than long (14:13) and 
rugose dorsally, becoming smooth at pos- 
terior margin. Metasoma. Ovate, about 
1.5X as long as wide, with brown margin 
laterally interrupted by yellow about mid- 
point making the yellow area appear as an 
inverted "T" shape, ventrally light yellow 
to white, becoming dark yellow posteri- 
orly. Legs. Ratio of hind tibial spur l:spur 
2:tarsus 1:T2:T3:T4. 35:28:20:17:11:19. Fore- 
wing. Hyaline, about 2.2 X as long as 
wide. Costal cell with 1 and occasionally 
a partial second irregular row of setae 
ventrally. Venation yellow to white, ratio 
of postmarginal: stigmal (29:20). 

Male. — Unknown. 

Hosts. — Motya abseuzalis (Noctuidae). 

Distribution. — Known only from the 
type locality. 

Types. — Holotype female: Costa Rica, 
Guanacaste Pro v., Ai'ea de Conservacion 
Guanacaste, Lambert N309250 E353000, 
2m., VII. 6, 1995, 95-SRNP-6049, D.H. Jan- 
zen & W. Hallwachs. ex. Motya abseuzalis 
(deposited in INBIO). Paratypes: 6 females 
with the same data as holotype (deposited 
in USNM and BMNH). 

Etymology. — This species is named in 
honor of Irene Carrillo Carillo in special 
recognition of her dedicated attention to 
the dining operations in Sector Santa Rosa 
of the Area de Conservacion Guanacaste. 

EupJectrus ivonae Schauff, new species 

(Figs. 16, 32, 52, 53) 
Diagnosis. — Face below and between to- 
ruli yellow, extending laterally to near eye 



208 



Journal of Hymenoptera Research 



and ventrally around mouth and gena 
(Fig. 16); legs yellow; one pair of setae ss2 
between lateral ocelli; seta S5 absent; lon- 
gitudinal carina on mesoscutum nearly 
complete, midlobe without small setae; 
scutellum finely longitudinally striate re- 
ticulate; metanotum bordered anteriorly 
and medially by small alveoli, medially 
expanded into a triangular flange; propo- 
deum laterad of median carina nearly 
smooth (lightly reticulate) (Fig. 53); dorsal 
metasoma with large central yellow area 
extending from just behind petiole poste- 
riorly for about 2/3 length, interrupted 
posteriorly by a central dark spot and be- 
coniing lighter brown again in posterior 
1/4, laterally dark brown. Male. Face with 
white area almost touching eye laterally; 
legs white; antenna with scape white, 
slightly swollen on ventral surface, with 
sensory area slightly darker and with sev- 
eral irregular rows of sensillae extending 
for about 3/4 length (Fig. 32). 

This species is similar to species like 
uiagdae, ronniei, and murine, which lack S5 
and have the face yellow. In this species, 
the face is more extensively yellow with 
the coloration extending laterad of the to- 
ruli over to and below the eyes. It does 
not, however, extend up the margin of the 
eyes as in flon/ne. 

Description. — Female. Body length 2.25- 
2.5 mm. Color: body mostly black except 
the following: face below and between to- 
ruli yellow, extending laterally to near eye 
and ventrally around mouth and gena 
(Fig. 16); antenna with scape white to light 
yellow, flagellum yellow to light brown; 
mandibles yellow; legs yellow; dorsal me- 
tasoma with large central yellow area ex- 
tending from just behind petiole posteri- 
orly for about 2/3 length, interrupted pos- 
teriorly by a central dark spot and becom- 
ing lighter brown again in posterior 1/4, 
laterally dark brown; ventral metasoma 
yellow. Head. Dorsally with one pair of 
minor seta ss2 inserted near occipital ca- 
rina between posterior ocelli (as in Fig. 
70), seta S5 absent, setal row sr2 present 



as 2-3 irregular rows of 12-18 setae reach- 
ing the bottom of the eye; occipital carina 
present medially; width of eye: width of 
face 12:38, posterior margin of eye sepa- 
rated from margin of head ventrally. Ratio 
of MS:EH 18:30; lateral ocellus more than 
1 diameter from eye (OD:OOD 13:10). 
Face below eyes rounded, not abruptly 
narrowing. Vertex under anterior ocellus 
reticulate. Toruli separated by about 2x 
their own diameters. Malar suture absent. 
Area under eye irregularly reticulate to 
alutaceous. Scape 6x as long as wide. Ra- 
tio of funicular segments 13:13:14:14:18, 
width 7 at Fl to 8 at F4. Mesosoiun. Pron- 
otum anterior to transverse carina with 
scattered setae, finely rugosely reticulate, 
posterior of carina more openly reticulate 
and shiny. Mesoscutum (Fig. 52) rugose 
reticulate in anterior 1/2, becoming more 
sniooth and shiny posteriorly with slight 
reticulation. Midlobe with median carina 
fading only at extreme anterior margin, 
otherwise complete, slightly sunken pos- 
teriorly, with no small setae antero-later- 
ally, posterior setae raised above surface 
on small tubercle. Dorsal axillar/scutellar 
n^iargin with broad, nearly straight deep 
furrow with narrow but flat bottom. Ax- 
illae shiny, openly reticulate, becoming 
smooth at posterior margin. Scutellum 
finely longitudinally striate reticulate, 
pointed anteriorly at axillar margin. Me- 
tanotum bordered anteriorly and medially 
by small alveoli, medially expanded into 
a triangular flange (Fig. 53) with median 
carina below. Fropodeum laterad of me- 
dian carina nearly smooth (lightly reticu- 
late) laterally to the step-like plica, median 
carina with anterior cup-like flange round- 
ed and deeply invaginated. Area around 
spiracle finely reticulate, lateral edge of 
spiracle raised above surface, with antero- 
lateral flange large and well defined, with 
9-10 setae laterad and below spiracle. Pos- 
terior margin of propodeum without ir- 
regular alveolae and carinae. Petiole in 
dorsal view wider than long (17:10) and 
rugose dorsally becoming smooth at pos- 



Volume 10, Number 2, 2001 



209 



terior margin. Mctasoma. Ovate, about 
1.5X as long as wide, with brown niargin 
laterally interrupted by yellow about mid- 
point making the yellow area appear as an 
inverted "T"shape. Legs. Ratio of hind tib- 
ial spur l:spur 2:tarsus 1:T2:T3:T4. 41:30: 
30:17:7:17. Forewing. Hyaline, about 2.2x 
as long as wide. Costal cell with 2 irreg- 
ular rows of setae ventrally. Venation yel- 
low to white, ratio of postmarginal: stig- 
mal (27:18). 

Male. — Similar to female except: body 
length 1.9 mm. Face with white area al- 
most touching eye laterally; legs white; 
dorsal metasoma with large central white 
spot, dark brown posteriorly; antenna 
with scape white, slightly swollen on ven- 
tral surface, with sensory area slightly 
darker and with several irregular rows of 
sensillae extending for about 3/4 length 
(Fig. 32); funicle ratios 13:13:14:13:18, 
width about 6 anteriorly to 7 posteriorly, 
with numerous semierect brown setae on 
each flagellomere. 

Hosts. — EiiscirrJioptcnis pmci/i (Noctui- 
dae). 

Distribiitio}!. — Known only from the 
type locality. 

Types. — Holotype female: Costa Rica, 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N313800 E359800, 
300m., V. 21, 1994, 94-SRNP-1656, D.H. 
Janzen & W. Hallwachs. ex. Eiiscirrlioptc- 
rus pocyi (deposited in INBIO). Paratypes: 
6 females and 1 male with same data as 
holotype (deposited in USNM). 

Etymology. — This species is named in 
honor of Ivon Traha Medrano in special 
recognition c^f her dedicated attention to 
the dining operations in Sector Santa Rosa 
of the Area de Conservacion Guanacaste. 

Euplcctrtis josei Schauff, new species 

(Figs. 35, 54-57) 

Diagnosis. — Face below and between to- 
ruli yellow; dorsally with two pairs of mi- 
nor seta ss2 between posterior ocelli (Fig. 
54); posterior margin of eye nearly contig- 
uous with posterior margin of head over 



most of length. Male antenna with scape 
white, slightly swollen (Fig. 57), with nar- 
row, elongate sensory area containing 2 ir- 
regular rows of sensillae extending almost 
entire length (Fig. 35). 

This species is recognizable by two 
characters: 2 pairs of setae between the lat- 
eral ocelli (ss2) and the posterior margin 
of the eye contiguous with the posterior 
margin of the head. Euplectrus floryac also 
has two pairs of ss2 setae, but the yellow 
color on the face runs to and partially up 
the margin of the eyes whereas in josci it 
is restricted to between and below the to- 
ruli. 

Description. — Female. Body length 2.2- 
2.5 mm. Color: body mostly black except 
the following: face below and between to- 
ruli yellow; antenna with scape white to 
light yellow, flagellum yellow; mandibles 
yellow; legs light yellow to white; dorsal 
metasoma with large central yellow to 
white area extending from just behind pet- 
iole posteriorly about 1/2-2/3 length of 
dorsum, becoming darker brown posteri- 
orly, lateral niargin brown anteriorly, then 
yellow medially and becoming dark 
brown posteriorly; ventral metasoma yel- 
low to white. Head. Dorsally with two 
pairs of minor seta ss2 between posterior 
ocelli, inserted adjacent to occipital carina 
(Fig. 54), all setae Sl-6 present, setal row 
sr2 present as 1-2 irregular rows of 12-15 
setae reaching the bottom of the eye; oc- 
cipital carina weak medially; width of eye: 
width of face: 35:15, posterior margin eye 
of nearly contiguous with posterior mar- 
gin of head over most of length. Ratio of 
MS:EH 13:33; lateral ocellus less than 1 di- 
ameter from eye (OD:OOD 8:5). Face be- 
low eyes rounded, not abruptly narrow- 
ing. Vertex under anterior ocellus reticu- 
late to alutaceous. Toruli separated by 
about 2X their own diameters. Malar su- 
ture absent below eye. Area under eye 
lightly reticulate to alutaceous. Scape 4x 
as long as wide. Ratio of funicular seg- 
ments 13:13:13:12:19, width 7 at Fl to 8 at 
club, flagellar segments without small 



210 



Journal of Hymenoptera Research 



whorls of brown setae basally. Mesosonw. 
Pronotum anterior to transverse carina 
with scattered setae, finely rugosely retic- 
ulate, posterior to carina more openly re- 
ticulate and shiny. Mesoscutum (Fig. 55) 
reticulate, becoming more smooth, and 
shiny posteriorly. Midlobe with median 
carina fading over anterior 1/4, otherwise 
complete, slightly sunken posteriorly, 
with no small setae antero-laterally, pos- 
terior setae even with surface. Dorsal ax- 
illar/scutellar margin with broad, nearly 
straight deep furrow with flat bottom. Ax- 
illae shiny, openly reticulate. Scutellum re- 
ticulate to alutaceous, smooth along pos- 
terior margin, slightly pointed at anterior 
margin with axillae. Metanotum bordered 
anteriorly by narrow band of small alve- 
oli, medially shiny and lightly reticulate 
with anterior edge slightly projected out- 
ward. Propodeum laterad of median cari- 
na shiny and sniooth to very lightly retic- 
ulate to the step-like plica, median carina 
with anterior cup-like flange nearly trian- 
gular, slightly invaginated. Area around 
spiracle reticulate, openly reticulate later- 
ally, spiracle slightly raised above surface 
and slanted so that opening is at a slight 
angle to the surface (Fig. 56), with antero- 
lateral flange present and obvious, with 8- 
12 setae laterad and below spiracle. Pos- 
terior margin of propodeum with a deep 
alveolus at posterior margin adjacent to 
plica. Petiole in dorsal view slightly longer 
than wide (11:10), rugose dorsally with ir- 
regular longitudinal carina and smooth at 
posterior margin. Metasoina. Ovate, about 
1.5-2X as long as wide. Legs. Ratio of 
hind tibial spur l:spur 2:tarsus 1:T2:T3:T4. 
34:25:23:16:10:15. Forewing. Hyaline, 
about 2.3 X as long as wide. Costal cell 
with 2 irregular rows of setae ventrally. 
Venation yellow to white, ratio of post- 
marginal: stigmal (30:18). 

Male. — Similar to female except: Body 
length 2.25 mm. Antenna with scape 
white, (Figs. 35, 57) slightly swollen, with 
narrow, elongate sensory area containing 
2 irregular rows of sensillae extending al- 



most entire length; funicle ratios 14:14:14: 
14:20 width 6-7, with scattered or whorled 
semierect brown setae on each flagelloni- 
ere. 

Hosts. — Pacctcs luiiodcs (Noctuidae). 

Distribution. — Known only from the 
ACG. 

Types. — Holotype female: Costa Rica, 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N314800 E360500, 
300m., VII. 8, 1993, 93-SRNP-3064, D.H. 
Janzen & W. Hallwachs. ex. Pnectes luuodes 
(deposited in INBIO). Paratypes: 2 males 
with the same data as holotype; other 
specimens with some of the same infor- 
mation as holotype are except: 2 females 
with Lambert N313800 E359800, 300m., 
VII. 17. 1984, 84-SRNP-1504; 2 males and 

1 female with Lambert N313800 E359800, 
300m., VII. 18. 1984, 84-SRNP-1433; 2 fe- 
males with Lambert N314800 E360500, 
300m., VII. 6. 1993, 93-SRNP-2869; Imale 
and 1 female with Lambert N313800 
E359800, 300m., VII. 4. 1993, 93-SRNP- 
2871; 1 male with Lambert N314800 
E360500, 300m., VII.7.1993, 93-SRNP-3093; 

2 males and one female with Lambert 
N314400 E358900, 280m., VII.4.1995, 95- 
SRNP-6055 (deposited in USNM, BMNH, 
and CNC). 

Eti/iuology. — This species is named in 
honor of Jose Eras Pineda in special rec- 
ognition of his dedicated management of 
the dining operations in Sector Santa Rosa 
of the Area de Conservacion Guanacaste. 

Ettplectrus magdae Schauff, new species 

(Figs. 33, 58-60) 

Diagnosis. — Face below and between to- 
ruli yellowish brown (as in Figs. 13, 14); 
legs light yellow to white; one pair of 
small setae (ss2) between lateral ocelli (as 
in Fig. 54); all ntajor setae SI -6 present; 
metanotum bordered anteriorly by a large 
nearly continuous invagination sometimes 
divided medially into two alveoli (Fig. 59); 
propodeum laterad of median carina retic- 
ulate, median carina with anterior cup-like 
flange nearly triangular and only slightly 



Volume 10, Number 2, 2001 



211 



invaginated; petiole wider than long. Male 
antenna with scape white, swollen on lat- 
eral surface (Fig. 60), with sensory area in- 
vaginated and with 3-4 irregular rows of 
sensillae extending nearly entire length 
(Fig. 33). 

This species is similar to E. nunc which 
shares the yellow face, yellow legs, and 
presence of all 6 pairs of large facial setae. 
In E. nunc the petiole is longer than wide 
while in E. uingdne the petiole is slightly 
wider than long. The male scape of E. uing- 
dne is more swollen and sunken with a 
larger sensory area than in £. nunc (Fig. 
34). 

Description. — Female. Body length 2.2- 
2.6 mm. Color: body mostly black except 
the following: face below and between to- 
ruli yellowish brown; antenna with scape 
white to light yellow, flagellum yellow to 
light brown; mandibles yellow to white; 
legs light yellow to white; dorsal metaso- 
ma with large central yellow area extend- 
ing from just behind petiole posteriorly for 
about 1/2 length, roughly hourglass 
shaped posteriorly dark brown to black, 
ventral metasoma yellow in anterior half, 
brown posteriorly. Hend. Dorsally with 
one pair of minor seta ss2 between pos- 
terior ocelli (as in Fig. 70), inserted dis- 
tinctly above occipital carina, all setae Sl- 
6 present, setal row sr2 present as 2-3 ir- 
regular rows of 20-25 setae reaching the 
bottom of the eye; occipital carina present 
medially; width of eye: width of face 14: 
40, posterior margin of eye separated from 
margin of head ventrally. Ratio of MS:EH 
16:30; lateral ocellus more than 1 diameter 
from eye (OD:OOD 14:10). Face below 
eyes rounded, not abruptly narrowing. 
Vertex under anterior ocellus smooth to 
lightly reticulate. Toruli separated by 
about 2X their own diameters. Malar su- 
ture absent. Area under eye smooth. Scape 
4X as long as wide. Ratio of funicular seg- 
ments 14:14:14:14:19, width 6 at Fl to 7 at 
club, each flagellar segment with 1-2 ir- 
regular whorls of brown setae basally. Mc- 
sosomn. Pronotum anterior to transverse 



carina with scattered setae, finely rugosely 
reticulate, posterior to carina niore openly 
reticulate and shiny. Mesoscutum (Fig. 58) 
reticulate, becoming more smooth, and 
shiny posteriorly. Midlobe with median 
carina fading only at extreme anterior 
margin, otherwise complete, slightly 
sunken posteriorly, with no small setae 
antero-laterally, posterior setae even with 
surface. Dorsal axillar/scutellar margin 
with broad, curved deep furrow with flat 
bottom. Axillae shiny, openly reticulate. 
Scutellum finely reticulate to alutaceous, 
shiny, more nearly rounded anteriorly at 
margin with axillae. Metanotum bordered 
anteriorly by a large nearly continuous in- 
vagination sometimes divided medially 
into two alveoli, medially shiny and light- 
ly reticulate, without median carina be- 
low. Propodeum (Fig. 59) laterad of me- 
dian carina reticulate to the step-like plica, 
median carina with anterior cup-like 
flange nearly triangular and only slightly 
invaginated. Area around spiracle finely 
reticulate, lateral edge of spiracle raised 
above surface, with antero-lateral flange 
reduced, with 7-8 setae laterad and below 
spiracle. Posterior margin of propodeum 
without irregular alveolae and carinae. 
Petiole in dorsal view slightly wider than 
long (15:12) anci rugose dorsally. Mctnso- 
mn. Ovate, about 1.7-2X as long as wide. 
Legs. Ratio of hind tibial spur l:spur 2: 
tarsus 1:T2:T3:T4. 40:30:27:16:11:18. Fore- 
wing. Hyaline, about 2.5 X as long as 
wide. Costal cell with 2 irregular rows of 
setae ventrally. Venation yellow to white, 
ratio of postmarginal: stigmal (33:17). 

Mnk. — Similar to female except: body 
length 1.9 mm. Face with white area 
slightly broader; legs white with distal 
femora, tibiae, and tarsi sometimes yel- 
low; dorsal metasoma with large central 
white spot, dark brown posteriorly; anten- 
na with scape white, (Figs. 33, 60) swollen 
on ventral surface, with sensory area in- 
vaginated and with 3^ irregular rows of 
sensillae extending nearly entire length; 
funicle ratios 13:13:14:13:18, width about 6 



212 



Journal of Hymenoptera Research 




Figs. 56-63. Eiiplcctyiis .scanning electron microgmpl-is. 56-57, E. josci. 56, Prdpodeiim. 57, Male scapes. 58- 
60, £. iiiiP^daL'. 58, Dorsal mesosoma. 5'), rrt>podeuni. ()(), Male scapes. 61-63, E. iiiiiriiu'. 61, Dorsal mesosoma. 
62, Male head and scape. 63, Closeup of male scape. 



VoLUMB 10, Number 2, 2001 



213 




Figs. 64-71. Euph'ctrus scanning electron micrograpiis. 64-67, £. orins. 64, Dorsal mesosoma. 65, Propodeum. 
66, Head, dorsal view. 67, Head, lateral view. 68-69, £. rojasi. 68, Dorsal mesosoma. 69, Propodeum, lateral 
view. 70-71, E. nviiiici. 70, Head and mesosoma. 71, Dorsal mesosoma. 



214 



Journal of Hymenoptera Research 



anteriorly to 7 posteriorly, with whorled 
semierect brown setae on base of each fla- 
gellomere. 

Hosts. — Dasylophia maxtla, D. basitinc- 
ta, D. nr. goraxa (all Notodontidae). 

Distribution. — Known only from the 
ACG. 

Ti/pes. — Holotype female: Costa Rica 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N318600 E375150, 
560m., IX. 12. 1995, 95-SRNP-9001, ex. Das- 
ylophia maxtla, D.H. Janzen & W. Hall- 
wachs (deposited in INBIO). Paratypes: 5 
males and 25 females with same data as 
holotype (deposited in USNM, BMNH, 
CNC). 

Otlier specimens examined. — All from the 
ACG, 4 specimens 94-SRNP-6167, ex. Chli- 
ara croesus; 4 specimens 96-SRNP-11096; 
1 specimen 93-SRNP-2905, ex. Dasylophia 
basitincta, 4 specimens 87-SRNP-1302, ex. 
Dasylophia not basitincta. 

Eti/Diologif. — This species is named in 
honor of Maria Magdalena Rodriguez 
Rodriguez in special recognition of her 
dedicated management of the main Ad- 
ministrative Office of the Area de Conser- 
vacion Guanacaste. 

Eiiplectnis marine Schauff, new species 

(Figs. 30, 61-63) 

Diagnosis. — Face below and between to- 
ruli yellow, extending slightly laterad of 
toruli, but not reaching eye (as in Fig. 13); 
one pair of setae ss2 between posterior 
ocelli, seta S5 absent (see Fig. 23); mesos- 
cutal midlobe without small setae anteri- 
orly; propodeum (Fig. 61) lightly reticu- 
late, often with irregular carinae laterad of 
median carina and appearing somewhat 
rugose; petiole in dorsal view wider than 
long. Male antenna with scape white, 
slightly swollen, with brown, elongate 
narrowly ovate sensory area containing 2- 
3 irregular rows of sensillae extending 
about 2/3 length (Fig. 30). 

This species is very similar to £. ronniei 
which also has a yellow face and lacks seta 
S5. EupJectrus marine can be distinguished 



from ronniei by the petiole which is as 
wide as long in that species, but wider 
than long in magdae. The niale scape of 
ronniei has the small brown sensory area 
much shorter (Fig. 31) and with only a sin- 
gle row of sensillae while in marine the 
sensory area is long and with at least two 
irregular rows of sensillae (Figs. 30, 62, 
63). 

Description. — Female. Body length 2.1- 
2.3 mm. Color: body mostly black except 
the following: face below and between to- 
ruli yellow, extending slightly laterad of 
toruli but not reaching eye; antenna with 
scape white to light yellow, flagellum yel- 
low or light brown; mandibles yellow to 
white; legs light yellow to white; dorsal 
metasoma with large central yellow to 
white area extending from just behind pet- 
iole posteriorly about 2/3 to 3/4 length of 
dorsum, becoming slightly darker poste- 
riorly, lateral brown margin continuous 
over length, ventral metasoma yellow to 
white. Head. Dorsally with one pair of mi- 
nor seta ss2 between posterior ocelli, seta 
S5 absent, setal row sr2 present as 1-3 ir- 
regular rows of 20-25 setae reaching the 
bottom of the eye; occipital carina fading 
medially; width of eye: width of face 11: 
33, posterior margin eye of nearly contig- 
uous with posterior margin of head most 
of length; ratio of MS:EH 15:30; lateral 
ocellus more than 1 diameter from eye 
(OD:OOD 6:8). Face below eyes rounded, 
not abruptly narrowing. Vertex under an- 
terior ocellus reticulate. Toruli separated 
by about 2x their own diameters. Malar 
suture absent. Area under eye lightly re- 
ticulate. Scape 6x as long as wide. Ratio 
of funicular segments 12:12:12:13:17, 
width 6 at Fl to 7 at club, flagellar seg- 
ments without small whorls of brown se- 
tae basally. Mesosoma. Pronotum anterior 
to transverse carina with scattered setae, 
finely rugosely reticulate, posterior to ca- 
rina more openly reticulate and shiny. Me- 
soscutum (Fig. 61) reticulate, becoming 
more smooth, and shiny posteriorly. Mid- 
lobe with median carina fading over an- 



VoLUMF 10, Number 1, 2001 



215 



terior 1/4, otherwise complete, slightly 
sunken posteriorly, with no small setae 
antero-laterally, posterior setae even with 
surface or slightly raised. Dorsal axillar/ 
scutellar margin with broad, nearly 
straight deep furrow with narrow, but flat 
bottom. Axillae shiny, openly reticulate. 
Scutellum shiny and lightly reticulate to 
alutaceous, pointed at anterior margin. 
Metanotum bordered anteriorly by a nar- 
row banci of small alveoli, medially shiny 
and lightly reticulate. Propodeum laterad 
of median carina shiny and openly retic- 
ulate to the step-like plica, median carina 
with anterior cup-like flange rounded and 
invaginated. Area around spiracle granu- 
late medially, openly reticulate laterally, 
spiracle slightly raised above surface par- 
allel to the surface of the propodeum, with 
antero-lateral flange large and obvious, 
with 8-9 setae laterad and below spiracle. 
Petiole in dorsal wider than long (20:15) 
and rugose dorsally with irregular longi- 
tudinal carina and smooth at posterior 
margin. Metasomn. Ovate, about 1.5-2X as 
long as wide. Legs. Ratio of hind tibial 
spur l:spur 2:tarsus 1:T2:T3:T4. 32:25:18: 
14:8:20. Forewing. Hyaline, about 2.3 X as 
long as wide. Costal cell with 2 irregular 
rows of setae ventrally. Venation yellow 
to white, ratio of postmarginal: stigmal 
(32:18). 

Male. — Similar to female except: Body 
length 1.7-1.9 mm. Face yellow to white, 
legs white to light yellow, metasoma with 
central white spot extending only about 
1/2 to 2/3 of length, dark brown posteri- 
orly, laterally dark brown to light brown. 
Antenna (Figs. 30, 62, 63) with scape 
white, slightly swollen, with brown, elon- 
gate narrowly ovate sensory area contain- 
ing 2-3 irregular rows of sensillae extend- 
ing about 2/3 length (Fig. 30); funicle ra- 
tios 11:10:11:11:16, width 6-7, without 
scattered or whorled semierect brown se- 
tae on each flagellomere. 

Hosts. — Coucaua Mumiissiifui (Noctui- 
dae), Elyiiiiotis attcmiatti (Notodontidae), 
Dnsi/lophia nr. gornxn (Notodc^ntidae). 



Distribution. — Known only from Guan- 
acaste. 

Ti/pcs. — Holotype female: Costa Rica, 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N309700 E352300, 
5m., V. 18, 1994, 94-SRNP-1063, D.H. Jan- 
zen & W. Hallwachs. ex. Concana mun- 
dissima (deposited in INBIO). Paratypes: 

2 females and 1 male all with the same 
data as holotype; other specimens with 
some of the same information are except: 

3 males and 1 female with Lambert 
N316100 E360300, 310m., VI. 13. 1993, 93- 
SRNP-1585; 1 male and 3 females with 
Lambert N318500 E359850, 240m., V. 24. 
1992, 92-SRNP-3032; 1 female with Lam- 
bert N309700 E352300, 5m., V. 19. 1994, 
94-SRNP-1083; 2 males with Lambert 
N309700 E352300, 5m., V. 16. 1994, 94- 
SRNP-1075; 1 female with Lambert 
N309700 E352300, 5m., V. 16. 1994, 94- 
SRNP-1071; 1 female with Lambert 
N316100 E360300, 310m., VI.15.1993, 93- 
SRNP-1577; 2 males with Lambert 
N309700 E352300, 5m., V. 19. 1994, 94- 
SRNP-1217; 1 female with Lambert 
N316100 E360300, 300m., Vl.14.1993, 93- 
SRNP-1583; 1 female with Lambert 
N309700 E352300, 5m., V. 16. 1994, 94- 
SRNP-1088; 1 male with Lambert N309700 
E352300, 5m., V.24.1994, 94-SRNP-1216; 1 
male and 1 female with Lambert N309700 
E352300, 5m., V.15.1994, 94-SRNP-1055; 1 
male and 1 female with Lambert N309700 
E352300, 5m., V.19.1994, 94-SRNP-1160; 1 
male with Lambert N309700 E352300, 5m., 
V.18.1994, 94-SRNP-1081; 1 male and 1 fe- 
male with Lambert N309700 E352300, 5m., 
V. 14. 1994, 94-SRNP-1102; 1 female with 
Lambert N318500 E359850, 240m., 
VII.22.1992, 92-SRNP-3141; 1 male with 
Lambert N318500 E359850, 240m., 
Vn.19.1992, 92-SRNP-2923; 1 female with 
Lambert N3 18500 E359850, 240m., 
VII.22.1992, 92-SRNP-3142; 1 male with 
Lambert N309700 E352300, 5m., V.18.1994, 
94-SRNP-1159; 1 female with Lambert 
N309700 E352300, 5m., V. 16. 1994, 94- 
SRNP-1090; 1 male, 2 females with Lam- 



216 



Journal of Hymenoptera Research 



bert N31N500 E359850, 240m., VII.19.1992, 
92-SRNP-2924; 1 female with Lambert 
N312150 E357200, 250m., VII.24.1992, 92- 
SRNP-3297; 1 male with Lambert N309700 
E352300, 5m., V.18.1994, 94-SRNP-1093; 1 
female with Lambert N318500 E359850, 
240m., Vll.19.1992, 92-SRNP-2924; 1 male 
and 1 female with Lambert N314800 
E360500, 300m., VII. 27.1993, 93-SRNP- 
3731; 1 male and 1 female with Lambert 
N314800 E360500, 300m., VII.23.1992, 92- 
SRNP-4426; 1 male with Lambert N314800 
E360500, 300m., VII. 23. 1994, 94-SRNP- 
5595; 1 female with Lambert N309450 
E355300, 10m., VI.5.1992, 93-SRNP-993; 2 
males and 1 female with Lambert N308900 
E355700, 10m., V.24.1996, 96-SRNP-1314; 1 
female with Lambert N316100 E360300, 
310m., VI. 14.1993, 93-SRNP-1582; 1 male 
and 1 female with Lambert N314800 
E360500, 300m., VIL19.1994, 94-SRNP- 
5588; 1 male with Lambert N314800 
E360500, 300m., VII. 31. 1991, 91-SRNP- 
2245 (paratypes deposited in USNM, IN- 
BIO, BMNH, and CNC). 

Other spicciiiiciis exniiiiiied. — 3 specimens 
92-SRNP-4431, ex. Eh/miotis sp.; 2 speci- 
mens 91-SRNP-1091, ex. Coiictiiui imiiidis- 
sifiia. 

Eti/iuologi/. — This species is nan"ied in 
honor of Maria De Los Angeles Guevara 
Rojas in special recognition of her dedi- 
cated attention to the dining operations in 
Sector Santa Rosa of the Area de Conser- 
vacion Guanacaste. 

Euplectrtis orias Schauff, new species 

(Figs. 15, 37, 64-67) 

Ding)iosis. — Female face beiovv toruli 
dark brown (Fig. 15), legs with midcoxa 
brown and hind coxa brown to black, hind 
feniur brown distally; antenna with scape 
yellow, flagellum yellow or light brown; 
mandibles yellow; with one pair of minor 
seta ss2 between posterior ocelli (Fig. 66), 
malar suture present below eye (Fig. 67); 
petiole as wide as long. 

This is one of a small group of species 



with a dark face and dark hind coxae (~(?- 
iiionii, valvcrdei and rojnsi). It can be differ- 
entiated from zamorai, rojasi, and valverdci 
by the presence of a single pair of ss2 setae 
between the lateral ocelli (setae absent in 
zamorai, rojasi, and valvcrdei). In addition, 
in zamorai the antennal tlagellum has the 
last two segments distinctly darker than 
the preceding segments (segments only 
gradually becoming darker in orias). In £. 
xiomarae, which also has a dark face and 
hind coxae, the malar suture is absent (this 
suture can be difficult to assess since it is 
often weakly expressed so caution is 
urged). 

Description. — Female. Body length 1.6- 
1.7 mm. Color: body mostly black except 
the following: face below and between to- 
ruli brown (Fig. 15); antenna with scape 
yellow, flagelluni yellow or light brown; 
mandibles yellow; legs yellow-brown ex- 
cept midcoxa brown, hind coxa mostly 
dark brown to black, hind femur with dis- 
tal half brown; dorsal metasoma with 
large central yellow area extending from 
just behind petiole posteriorly about 1/2 
length of dorsum, beconiing darker brown 
posteriorly, lateral margin brown, ventral 
metasoma yellow in anterior half, then 
dark brown. Head. Dorsally with one pair 
of minor seta ss2 between posterior ocelli, 
inserted adjacent to occipital carina (Fig. 
66), all setae Sl-6 present, setal row sr2 
present as 1-2 irregular rows of 8-10 setae 
reaching the bottom of the eye; occipital 
carina weak niedially; width of eye: width 
of face: 12:39, posterior margin eye of not 
nearly contiguous with posterior margin 
of head over most of length; ratio of MS: 
EH 12:24; lateral ocellus more than 1 di- 
ameter from eye (OD:OOD 8:12). Face be- 
low eyes rounded, not abruptly narrow- 
ing. Vertex under anterior ocellus reticu- 
late. Toruli separated by about 2.5 X their 
own dianieters. Malar suture present be- 
low eye, fading ventrally. Area under eye 
lightly reticulate to alutaceous. Scape 4X 
as long as wide. Ratio of funicular seg- 
ments 8:7:7:8:12, width 5 at Fl to 6 at club. 



Volume 10, Number 2, 2001 



217 



flagellar segments with small whorls of 
brown setae basally. Mesosoiiin. Pronotum 
anterior to transverse carina with scattered 
setae, finely rugosely reticulate, posterior 
to carina more openly reticulate and shiny. 
Mesoscutum (Fig. 64) reticulate, becoming 
more smooth, and shiny posteriorly. Mid- 
lobe with median carina well developed, 
fading over anterior 1/4, otherwise com- 
plete, not noticeably sunken posteriorly, 
with no small setae antero-laterally, pos- 
terior setae even with surface. Dorsal ax- 
illar/scutellar margin with broad, nearly 
straight deep furrow with flat bottom. Ax- 
illae shiny, openly reticulate. Scutellum 
shiny, lightly reticulate to alutaceous, 
smooth along posterior margin, slightly 
pointed at anterior margin with axillae. 
Metanotum borciered anteriorly by a nar- 
row band of small alveoli, medially shiny 
and lightly reticulate (Fig. 65). Propodeum 
laterad of median carina shiny and 
smooth to the step-like plica, median ca- 
rina with anterior cup-like flange rounded 
and invaginated. Area around spiracle 
granulate medially, openly reticulate lat- 
erally, spiracle slightly raised above sur- 
face and slanted so that opening is at a 
slight angle to the surface, with antero-lat- 
eral flange present, but somewhat re- 
duced, with 6-8 setae laterad and below 
spiracle. Petiole in dorsal as wide as long 
(12:12), rugose dorsally with irregular lon- 
gitudinal carina and smooth at posterior 
margin. Meiasoinn. Ovate, about 1.1-1.5X 
as long as wide. Legs. Ratio of hind tibial 
spur l:spur 2:tarsus 1:T2:T3:T4. 32:22:20: 
13:8:12. Forewing. Hyaline, about 2.3x as 
long as wide. Costal cell with 2 irregular 
rows of setae ventrally. Venation yellow 
to white, ratio of postmarginal: stigmal 
(16:12). 

Male. — Similar to female except: body 
length 1.6mm; antenna with scape white, 
slightly swollen, with narrow, elongate 
sensory area containing 2 irregular rows 
of sensillae extending about 3/4 length 
(Fig. 37); funicle ratios 7:7:7:7:11 width 5- 



6, with scattered or whorled semi-erect 
brown setae on each flagellomere. 

Hosts. — Unknown Geometridae. 

Distribution. — Widely ciistributed in 
Costa Rica. 

Types. — Holotype female: Costa Rica, 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N314800 E360500, 
300m., VI. 24, 1993, 93-SRNP-2241, D.H. 
Janzen & W. Hallwachs. ex. Geometridae 
(deposited in INBIO). Paratypes: 7 females 
and 1 male with the same data as holo- 
type; 1 female Costa Rica: Alajuela, 5km 
W. San Ramon, 1200m, IV. 1997, O. Castro 
& P. Hanson; 2 females Costa Rica: Pun- 
tarenas, San Vito, Estac. Biol. Las Alturas, 
1500m, IV. 1992, Hanson & Godoy; 1 fe- 
male Costa Rica: Cartago, 4km NE Canon, 
Genesis II, 2300m, VIII. 1995, P. Hanson; 1 
female Costa Rica: San Jose, Zurqui de 
Moravia, 1600m, X-XII. 1990, col. P. Han- 
son; 1 female Costa Rica: San Jose, Cerro 
de la Muerte, 19km S,3W Empalme, 
2600m, IX. 1992, Hanson & Godoy; 1 fe- 
male Costa Rica: Puntarenas, 23km N. 
Puerto Jimenez, LaPalma, 10m, XI-XII. 
1992, col. P. Hanson (deposited in USNM 
with 1 female each to CNC and BMNH). 

Etymology. — This species is named in 
honor of Julio Diaz Orias in special rec- 
ognition of his niany years of steadfast 
and energetic management of the fire pre- 
vention and control program of the Area 
de Conservacion Guanacaste. 

Enplectnis rojasi Schauff, new species 

(Figs. 68, 69) 

Diagnosis. — Face below toruli dark yel- 
low, brown under and between toruli; 
mandibles yellow; hind coxa mostly black; 
posterior margin of scutellum overlapping 
anterior metanotum (Figs. 68, 69); anterior 
extension of median propodeal carina flat- 
tened, not cup-like; petiole 1.5X as long as 
wide. 

The flattened anterior cup-like flange of 
the median propodeal carina in this spe- 
cies is quite distinctive with anterior end 
of median carina usually expanded into a 



218 



Journal of Hymenoptkra Research 



rounded or triangular and invaginated 
"cup." In addition, this is the only speci- 
men I have examined where the face un- 
der the toruli is brown medially between 
the toruli and becomes yellow laterally 
under the toruli. Of the species treated 
here, the anterior edge of the metanotum 
being covered by the posterior margin of 
the scutellum is also unusual. However, in 
E. carlowac, the lateral margins of the scu- 
tellum project over the lateral edges of the 
metanotum and the senior author has seen 
other specimens of apparently undescri- 
bed species which also have the scutellum 
overhanging the metanotum. 

Description. — Female. Body length 2.2 
mm. Color: body mostly black except the 
following: antenna with scape yellow to 
brown, pedicle becoming dark brown api- 
cally, flagellum light brown becoming 
darker brown apically; mandibles yellow; 
enlarged setae on vertex yellowish brown; 
legs yellow except hind coxa mostly black; 
dorsal metasoma mostly dark brown to 
black with small yellow spot antero-me- 
dially, ventral metasoma dark brown be- 
hind petiole, then yellow to about mid- 
point, then dark brown. Head. Dorsally 
with 5 minor seta ss2 between posterior 
ocelli, all seta Sl-6 present, setal row sr2 
present as 2 irregular rows of 15-20 setae 
reaching to bottom of eye; occipital carina 
weak medially; width of eye: width of face 
(35:13), posterior margin eye of not nearly 
contiguous with posterior margin of head 
over most of length. Ratio of MS:EH 16:30; 
lateral ocellus more than 1 diameter from 
eye (OD:OOD 5:8). Face below eyes round- 
ed, not abruptly narrowing. Vertex under 
anterior ocellus reticulate to ahitaceous. 
Toruli separated by about 2x their own 
diameters. Malar suture absent. Area un- 
der eye lightly reticulate. Scape 6X as long 
as wide. Ratio of funicular segments 10:1 1: 
11:11:18, width 5 at Fl to 6 at club, flagel- 
lar segments with small whorls of brown 
setae basally. Mesosomii. Pronotum anteri- 
or to transverse carina with scattered se- 
tae, finely rugosely reticulate, posterior to 



carina reticulate and shiny. Mesoscutuni 
(Fig. 68) reticulate, becoming more 
smooth, and shiny posteriorly. Mid lobe 
with median carina fading in anterior half, 
not noticeably sunken posteriorly, with no 
small setae antero-laterally, posterior setae 
even with surface. Dorsal axillar/scutellar 
margin with narrowed, nearly straight 
deep furrow without flat bottom except 
medially. Axillae openly reticulate. Scutel- 
lum reticulate to alutaceous, smooth along 
posterior margin, posterior margin ex- 
tended over anterior margin of metano- 
tum, anterit^r niargin pointed. Metanotum 
anteriorly covered by scutellum, medially 
shiny and smooth to lightly reticulate (Fig. 
69). Propodeum laterad of median carina 
shiny and smooth to the step-like plica 
with occasional slight hints of reticulation, 
median carina with anterior cup-like 
flange flattened, not expanded and cup- 
like. Area around spiracle reticulate, spi- 
racle slightly raised above surface and tilt- 
ed toward median carina, with antero-lat- 
eral flange present, with 11 setae laterad 
and below spiracle. Petiole in dorsal view 
1.5X as long as wide (15:10), rugose dor- 
sally. Metasoma. Ovate, about 1.5x as long 
as wide. Legs. Ratio of hind tibial spur 1: 
spur 2:tarsus 1:T2:T3:T4. 40:27:30:16:10:16. 
Forewing. Hyaline, about 2.3 X as long as 
wide. Costal cell with 1-2 irregular rows 
of setae ventrally. Venation yellow light 
brown, ratio of postmarginal: stigmal (41: 
23). 

Male. — Unknown. 

Hosts. — Unknown. 

Distribution. — Known only from the 
type locality. 

Ti/pes. — ^Holotype female on point (an- 
tenna and wing slide-mounted) with data 
"Costa Rica, Cartago, 4 Km. NE., Cafion 
Genesis 11, 2350M. VI. 1995. P. Hanson." 
(deposited in USNM). 

Notes. — The presence of 5 setae ss2 is 
probably an anomalv- Other specimens 
rarely show 1 seta or 3 setae when the 
usual number is 0, 2, or 4. In this case, the 



Volume 10, Number 2, 2001 



219 



usual number for this species is probably 
4 setae in this location. 

Etymology. — This species is named in 
honor of Maria Zulay Guevara Rojas in 
special recognition of her dedicateci atten- 
tion to the Research Center and the dor- 
mitories in Sector Santa Rosa of the Area 
de Conservacion Guanacaste. 

Euplectriis ronniei Schauff, new species 

(Figs. 31, 70, 71, 72, 73) 

Diagnosis. — Face below and slightly lat- 
erad of toruli yellow; with one pair of se- 
tae ss2 between posterior ocelli (Fig. 70), 
seta S5 absent (see Fig. 23), scutellum 
shiny and lightly reticulate, petiole as 
wide as long. Male scape with small re- 
stricted brown spot on ventral surface, 
single short row of sensillae (Fig. 31). 

This species is similar to £. ninriac which 
has a similar overall appearance lacking 
the S5 setae and with a yellow face, but 
tends to be somewhat larger (generally 
over 2 mm) and the propodeum laterad of 
the median carina is nearly smooth where- 
as it is more irregularly rugose in luariae. 
Euplectriis ronniei is most easily diagnosed 
by the scape (Figs. 31, 73) of the males 
which have a small restricted brown patch 
on the antero-ventral surface with only a 
single small row of sensillae. 

Description. — Female. Body length 1.6-2 
mm. Color: body mostly black except the 
following: face below and between toruli 
yellow, extending laterally below toruli, 
but not reaching eye; antenna with scape 
whitish, flagellum yellow; mandibles yel- 
low; legs yellow to white; dorsal metaso- 
ma with large central yellow area extend- 
ing from just behind petiole posteriorly 
about 1/2 length of dorsum, becoming 
darker brown posteriorly, lateral margin 
brown, ventral metasoma yellow to white 
in anterior half, then becoming dark yel- 
low. Head. Dorsally with one pair of minor 
seta ss2 between posterior ocelli (Fig. 70), 
seta S5 absent, setal row sr2 present as 1- 
2 irregular rows of 13-15 setae reaching 
the bottom of the eye; occipital carina 



weak medially; width of eye: width of 
face: 12:35, posterior margin eye of not 
nearly contiguous with posterior margin 
of head over most of length; ratio of MS: 
EH 14:25; lateral ocellus more than 1 di- 
ameter from eye (OD:OOD 6:7). Face be- 
low eyes rounded, not abruptly narrow- 
ing. Vertex under anterior ocellus reticu- 
late. Toruli separated by about 2X their 
own diameters. Malar suture absent below 
eye. Area under eye lightly reticulate to 
alutaceous. Scape 5X as long as wide. Ra- 
tio of funicular segments 9:9:9:9:14, width 
5 at Fl to 6 at club, flagellar segments with 
small whorls of brown setae basally. Me- 
sosoina. Pronotum anterior to transverse 
carina with scattered setae, finely rugosely 
reticulate, posterior to carina more openly 
reticulate and shiny. Mesoscutum (Fig. 71) 
reticulate, becoming more smooth, and 
shiny posteriorly. Midlobe with median 
carina well developed, fading over ante- 
rior 1/3 to 1/4, otherwise complete, not 
noticeably sunken posteriorly, with no 
small setae antero-laterally, posterior setae 
even with surface. Dorsal axillar/scutellar 
margin with broad, nearly straight deep 
furrow with narrow, but flat bottom. Ax- 
illae shiny, openly reticulate. Scutellum 
shiny, lightly reticulate to alutaceous, 
smooth along posterior margin, pointed at 
axillar/scutellar margin. Metanotum bor- 
dered anteriorly by narrow band of small 
alveoli, medially shiny and lightly reticu- 
late. Propodeum (Fig. 72) lateraci of me- 
dian carina shiny and lightly reticulate to 
the step-like plica, median carina with an- 
terior cup-like flange rounded and inv^ag- 
inated. Area around spiracle dull, openly 
reticulate laterally, spiracle slightly raised 
above surface and slanted so that opening 
is at a slight angle to the surface, with an- 
tero-lateral flange present, but somewhat 
reduced, with 9-12 setae laterad and be- 
low spiracle. Petiole in dorsal view as 
wide as long (11:11), rugose dorsally with 
irregular longitudinal carina and almost 
smooth at posterior margin. Metasoma. 
Ovate, about 1.4-1.5X as long as wide. 



220 



Journal of Hymenoptera Research 



Legs. Ratio of hind tibial spur l:spur 2: 
tarsus 1:T2:T3:T4. 28:16:20:12:8:13. Fore- 
wing, i-iyaline, about 2.3 x as long as 
wide. Costal cell with 2 irregular rows of 
setae ventrally. Venation yellow to white, 
ratio of postmarginal: stigmal (22:16). 

MnJe. — Similar to female except: body 
length 1.6 mm. Face white to yellow; an- 
teiinal scape with small brown spot at api- 
cal ventral margin (Fig. 31) very slightly 
swollen, with narrow sensory area con- 
taining only 1 irregular row of sensillae 
extending about 1/4 length; funicle ratios 
9:10:10:10:17 width 5 near base, 6 at club, 
with few scattered or whorled semierect 
brown setae on each flagellomere; meta- 
soma white in dorsal 1/2 then dark 
brown, ventrally yellow becoming dark 
brown posteriorly. 

Hosts. — Oxidcrcin toxcn (Noctuidae), 
Cnutcthin spuria (Sphingidae). 

Dlstribiitio)!. — Known only from the 
ACG. 

Types. — Female holotype: Costa Rica 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N314800 E360500, 
300m., Vll.26.1992, 92-SRNP-3415, ex. Cim- 
tethin spuria, D.H. Janzen & W. Hallwachs 
(deposited in INBIO). Paratypes: 3 males 
same as holotype except with Lambert 
N313400 E358900, 280m., Vlll. 28.1995, 95- 
SRNP-8256, ex. Oxidcrcia toxca; 1 female 
with Lambert N314800 E360500, 300m., 
Vll. 26.1992, 92-SRNP-3415, ex. Cautcihia 
spuria (deposited in USNM) 

Etymology. — This species is named in 
honor of Ronald Hernandez D'Avanzo in 
recognition of his dedicated management 
of the Human Resources office of the Area 
de Conservacion Guanacaste. 

Eiiplcctrus solitariiis Ashmead 

Eiiplcctnis solitariiis Asiimecid 1904:517. 

Dia^^iiosis. — Face under toruli honey yel- 
low, lighter area restricted to just under 
toruli; all major seta 1-6 present (see Fig. 
23); one pair of small setae (ss2) between 
posterior ocelli (as in Fig. 70); first funicle 



2x as long as wide; legs yellow; scutellum 
reticulate; petiole longer than wide; me- 
tasoma ovate with small yellow spot an- 
terio-medially becoming brown then dark 
brown laterally and posteriorly. 

This species is very similar to E. couisto- 
cl<ii but can be differentiated by the petiole 
which is slightly wider than long (longer 
than wide in solitarius). In addition, the 
first funicle in solitarius is 3x as long as 
wide while in coiustockii Fl is 2x as long 
as wide. 

I have seen very few speciniens of this 
species and it is defined here mostly on 
the basis of the holotype which is missing 
most of its antennae. I have seen no males 
that I can definitively attribute to this spe- 
cies. 

Distribution. — Mexico south to Brazil 
and Ecuador 

Hosts. — Unknown. 

Types. — Holotype female deposited in 
USNM, type no. 60573. Erroneously la- 
belled as a male, this type is missing most 
of the antennae. 

Etipylectnis valverdei Schauff, new 
species 

(Figs. 17, 20, 21, 38, 74, 75, 76) 

Diagnosis. — Face below toruli dark 
brown or black (as in Fig. 15); mandibles 
brown; hind femur and hind coxa darker 
than other legs (honey yellow to brown); 
minor seta ss2 absent between posterior 
ocelli (as in Fig. 82); anterior metanotum 
with narrow line of alveoli; malar suture 
visible at least near eye; propodeum lat- 
erad of median carina smooth. Male scape 
yellow and greatly swollen (Figs. 17, 38), 
surface covered with large sensillae (Figs. 
20, 21, 75, 76). 

In contrast to some of the other species 
with dark faces and hind coxae, the man- 
dibles of this species are also brown (man- 
dibles yellow in xioinarae and orias). The 
mandibles of zaniorai are also brown, but 
that species has the antenna with the last 
two antennal flagellomeres dark contrast- 
ing sharply with the lighter preceding seg- 




Figb. 72-74. Liiplcctnis scanning electron iiiKiugiaphs. 72-73, £. ivmici. 72, Propodeuni. 73, Head, lateral 
view, male. 74-76, £. valvcnici. 74, Dorsal mesosoma. 75, Male antennae and head. 76, Male scape. 77-79, £. 
'umJtcii. 77, Dorsal mesosoma. 78. Propodeum. 79, Head. 



222 



JOURNAI. OF H^'MFNOPTHRA RESEARCH 




I i,t;s. ,S(I~,S7. Liiplectnis scanninj^ citeliuii iiii>.ioi;ici|.iiij,. oW-(S4, L. wuHcn. (SO, Head lateral view. 81, Head, 
ventro-lateral view. 82, Head, postero-dorsal view. 83, Head, frontal \iew. 84. Dorsal mesosoma. 85-87, E. 
xioniarnr. 85, Head, postero-dorsal view. 86, Dorsal mesosoma. 87, Thorax and propodeum. 



Volume 10, Number 2, 2001 



Z23 



ments. In addition, E. valvcnici has the ma- 
lar suture at least partially complete while 
in zaniorai and xiomarnc it is absent. Males 
of this species are very distinctive with 
greatly enlarged yellow scapes (Fig. 17) 
covered on all surfaces with distinctly 
granular sensillae (when viewed on slide) 
(Figs. 20, 21). 

Description. — Female. Body length 1.7- 
2.1 mm. Color: body niostly black except 
the following: antenna with scape white to 
yellow, flagellum yellow to light brown; 
mandibles brown; enlarged setae on ver- 
tex silver; legs white to yellow except hind 
coxa and distal half of hind femur honey 
yellow to brown; dorsal metasoma with 
large central inverted T-shaped yellow 
area extending from just behind petiole 
posteriorly about 1/2 length of dorsum, 
becoming darker brown posteriorly, lat- 
eral margin brown except medially, ven- 
tral metasoma yellow or white in anterior 
half, then dark brown. Head. Dorsally with 
minor seta ss2 absent between posterior 
ocelli, all setae Sl-6 present, setal row sr2 
present as 2-3 irregular rows of 15-20 se- 
tae reaching to bottom of eye; occipital ca- 
rina weak to absent; width of eye: width 
of face (40:13), posterior margin eye of not 
nearly contiguous with posterior margin 
of head over most of length. Ratio of MS: 
EH 18:28; lateral ocellus more than 1 di- 
anieter from eye (OD:OOD 6:8). Face be- 
low eyes abruptly narrowing. Vertex un- 
der anterior ocellus reticulate to aluta- 
ceous. Toruli separated by about 2.5 X 
their own diameters. Malar suture pre- 
sent, but irregular, niarked in sonie spec- 
imens by change in sculpture. Area under 
eye lightly reticulate to alutaceous. Scape 
5x as long as wide. Ratio of funicular seg- 
ments 11:11:11:11:18, width 5 at Fl to 6 at 
club, flagellar segments with sniall whorls 
of brown setae basally. Mesosoiun. Prono- 
tum anterior to transverse carina with 
scattered setae, finely rugosely reticulate, 
posterior to carina reticulate and shiny. 
Mesoscutum reticulate, becoming more 
smooth, and shiny posteriorly. Midlobe 



(Fig. 74) with median carina nearly com- 
plete, fading at extreme anterior margin, 
not noticeably sunken posteriorly, with no 
small setae antero-laterally, posterior setae 
even with surface. Dorsal axillar/scutellar 
margin with broad, nearly straight deep 
furrow without flat bottom. Axillae nearly 
smooth, shiny, with faint open reticula- 
tion, pointed at anterior margin with ax- 
illae. Scutellum shiny, very lightly aluta- 
ceous, smooth along posterior margin. 
Metanotum anteriorly with narrow but 
distinct band of small alveoli, medially 
shiny and smooth. Propodeum laterad of 
median carina shiny and smooth to the 
step-like plica with occasional faint hints 
of reticulation, median carina with ante- 
rior cup-like flange rounded and invagi- 
nated. Area around spiracle reticulate, spi- 
racle slightly raised above surface and 
parallel to the surface, with antero-lateral 
flange present, with 12-15 setae laterad 
and below spiracle. Posterior niargin of 
propodeum with deep alveolus at poste- 
rior margin. Petiole in dorsal view slightly 
longer than wide (15:12), rugose dorsally. 
Metasouin. Ovate, about 1.3-1.6X as long 
as wide. Legs. Ratio of hind tibial spur 1: 
spur 2:tarsus 1:T2:T3:T4. 38:29:28:15:10:14. 
Forewing. Infuscate light brown, about 
2.3 X as long as wide. Costal cell with 1 
irregular row of setae ventrally. Venation 
yellow light brown, ratio of postmarginal: 
stigmal (22:16). 

Male. — Similar to female except: scape 
yellow and greatly swollen (Figs. 17, 38, 
75, 76) and surface covered with large 
granulate appearing sensillae (Figs. 20, 
21); face very narrow below toruli; legs 
generally white except hind leg yellow; 12: 
13:14:14:20, width 4-6 with no noticeable 
brown setae. 

Hosts. — Unknown 

Distribution. — Known only from the 
type locality. 

Types. — Holotype female: Costa Rica, 
San Jose, Ciudad Colon, 800m, II. 1990, 
Col. Luis Fournier (deposited in INBIO). 
Paratypes: 4 females with same data as the 



224 



Journal of Hymenoptera Research 



holotype; 2 males and 1 female with same 
data as holotype except III-IV. 1990 (de- 
posited in USNM and BMNH). 

Eti/fiiologif. — This species is named in 
honor of Julio A. Quiros Valverde in rec- 
ognition of his many years of service and 
diligent management of police and protec- 
tion services for the Area de Conservacion 
Guanacaste. 

Euplectrus ivalteri Schauff, new species 

(Figs. 13, 29, 77-84, 89) 

Dingnosis. — First funicular segment 
nearly equal in length to club (similar to 
Fig. 25), face below toruli yellow, with 1 
pair of setae ss2 (as in Fig. 70) between 
posterior ocelli, toruli separated by about 
4x their own diameter; posterior margin 
of eye nearly contiguous with posterior 
margin of head over most of length (as in 
Fig. 80, 82); scutellum heavily sculptured 
(Fig. 84); postmarginal vein almost 2X 
stigmal (see Fig. 18). Male Fl subequal to 
club, club swollen with broad, ovate sen- 
sory area containing several irregular 
rows of sensillae extending about 3/4 
length (Fig. 30, 81). 

The widely separated toruli and long 
first funicular segment (almost as long as 
the club) niake this species easily recog- 
nizable among those with a yellow face. 
In addition, the scutellum is much more 
heavily sculptured than most of the other 
species. The antenna of £. hniisoiii is some- 
what similar with Fl almost the same 
length as the club, but in that species all 
the funicles are elongated and about 3x 
as long as wide. In addition, the eyes of 
waltcri are large and the hind margin is 
contiguous with the back of the head over 
most of its length. 

Description. — Female. Body length 2.9- 
3.2 mm. Color: body mostly black except 
the following: face below and between to- 
ruli yellow (Fig. 13); antenna with scape 
white to light yellow, flagellum yellow or 
light brown; mandibles yellow to white; 
legs light yellow to white; dorsal metaso- 
ma with large central yellow to white area 



extending from just behind petiole poste- 
riorly about 1/2 length of dorsum, becom- 
ing darker brown posteriorly, lateral 
brown margin broken medially, ventral 
metasoma yellow to white. Head. Dorsally 
with one pair of nuinor seta ss2 between 
posterior ocelli (Fig. 82), inserted adjacent 
to occipital carina, setae Sl-6 present, setal 
row sr2 present as 1-3 irregular rows of 
15-20 setae reaching the bottom of the eye; 
occipital carina complete; width of eye: 
width of face:18:48, posterior margin eye 
of nearly contiguous with posterior mar- 
gin of head of most of length; ratio of MS: 
EH 17:31; lateral ocellus more than 1 di- 
ameter from eye (OD:OOD 8:10). Face be- 
low eyes abruptly narrowing. Vertex be- 
low anterior ocellus alutaceous. Toruli 
separated by about 4X their own diame- 
ters (Fig. 13). Malar suture absent. Area 
under eye lightly reticulate. Scape 6X as 
long as wide. Ratio of funicular segments 
19:17:17:15:20, width 8 at Fl to 9 at club, 
flagellar segments without small whorls of 
brown setae basally. Mesosoma. Pronotum 
anterior to transverse carina with scat- 
tered setae, finely rugosely reticulate, pos- 
terior to carina more openly reticulate and 
shiny. Mesoscutun"i reticulate (Fig. 84), 
nearly rugose anteriorly, becoming more 
smooth, and shiny posteriorly. Midlobe 
with median carina well developed, fad- 
ing over anterior 1/4, otherwise complete, 
slightly sunken posteriorly, small setae an- 
tero-laterally usually absent, rarely with 2, 
posterior setae even with surface or slight- 
ly raised. Dorsal axillar/scutellar margin 
with broad, curved deep furrow with flat 
bottom. Axillae shiny, openly reticulate. 
Scutellum heavily reticulate to alutaceous, 
smoc^th along posterior margin, anteriorly 
distinctly pointed at axillar/scutellar mar- 
gin. Metanotum bordered anteriorly by 
narrow band of small alveoli, medially 
shiny and lightly reticulate. Propodeum 
(Fig. 78) laterad of median carina shiny 
and openly reticulate to the step-like plica, 
median carina with anterior cup-like 
flange rounded and invaginated. Area 



Volume 10, Number 2, 2001 



T^^ 



around spiracle granulate medially, open- 
ly reticulate laterally, spiracle slightly 
raised above surface and slanted so that 
opening is almost perpendicular to sur- 
face, with antero-lateral flange large and 
obvious, with 8-12 setae laterad and be- 
low spiracle. Posterior margin of propo- 
deum without deep alveolus at posterior 
margin. Petiole in dorsal view as wicie as 
long or very slightly longer than wide (11: 
12), rugose dorsally with irregular longi- 
tudinal carina and smooth at posterior 
margin. Mctasoinn. Ovate, about 1.1-1.5X 
as long as wide. Legs. Ratio of hind tibial 
spur l:spur 2:tarsus 1:T2:T3:T4. 55:37:35: 
20:11:21. Forewing. Hyaline, about 2.3 x as 
long as wide. Costal cell with 2 irregular 
rows of setae ventrally. Venation yellow 
to white, ratio of postmarginal: stigmal 
(42:22). 

Male. — Similar to female except: body 
length 1.7-1.9 mm. Face yellow to white, 
legs white to light yellow, nietasc^ma with 
central white spot only extending about 
1/2 length, ciark brown posteriorly, later- 
ally dark brown to light brown, ventrally 
posterior half dark brown; antenna with 
scape white, swollen medially (Figs. 80, 
81), with broad, ovate sensory area con- 
taining several irregular rows of sensillae 
extending about 3/4 length (Figs. 13, 29); 
funicle ratios 17:16:15:13:18, width 5-6, 
with scattered or whorled semierect 
brown setae on each flagellomere. 

Hosts. — Mmiduca haruesi, M. dilucida, M. 
florestiiii, M. Iniiu^iiiosa, M. nistica, Pcrigcvi- 
in iliis (all Sphingidae). 

Notes. — Some setal variation on the me- 
soscutum has been observed in specimens 
of this species. While this character is gen- 
erally very stable and no small setae are 
present, there are occasionally setae pre- 
sent antero-laterally near the anterior en- 
larged setae on the midlobe. 

Types. — Holotype feniale: Costa Rica, 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N312450 E359500, 
270m., Vlll. 13, 1995, 95-SRNP-7544, D.H. 
Janzen & W. Hallwachs. ex. Mniiduca flo- 



restau (cieposited in INBIO). Para types: 2 
females all with the same data as holo- 
type; other specimens with some of the 
same information are except: 3 males and 
1 female with Lambert N313100 E359900, 
250m., VII. 30. 1992, 92-SRNP-3174; 3 
males with Lambert N313100 E359900, 
250m., VII. 30. 1992, 92-SRNP-4114; 2 
males and 2 feniales with Lambert 
N313800 E359800, 300m., VI. 30. 1984, 84- 
SRNP-653, ex. Mmiduca dilucida; 1 male 
with Lambert N313800 E359800,300m., VI. 
23. 1993, 93-SRNP-2239 ; 1 female with 
Lambert N314800 E360500, 300m., 
VII. 1.1984, 84-SRNP-805, ex. Mauduca dd- 
ucida; 1 male and 2 females with Lambert 
N313100 E359900, 250m., VII.27.1992, 92- 
SRNP-3018, ex. Mauduca nistica; 1 female 
with Lambert N315500 E360200, 300m., 
VIII.2.1992, 92-SRNP-3811; 2 females with 
Lambert N313100 E359900, 500m., 
VIII. 1.1992, 92-SRNP-3708; 1 male with 
Lambert N317800 E362600, 300m., 
VI.28.1984, 84-SRNP-623, ex. Mauduca dd- 
ucida; 1 female with Lambert N314800 
E360500, 300m., VIII.16.1981, 81-SRNP- 
1141, ex. Perigouia ilus; 2 males and one 
female with Lambert N312450 E359300, 
270m., VI.27.1996, 96-SRNP-6830; 1 female 
with Lambert N315500 E360200, 300m., 
VII. 2. 1992, 92-SRNP-2262, ex. Mauduca 
lanuginosa; 2 females and 1 male with 
Lambert N313400 E358900, 280m., 
X.3.1996, 96-SRNP-10435; 1 female with 
Lambert N313800 E359800, 300m., 
XII. 11. 1990, 90-SRNP-2507, ex. Mauduca 
rustica; 1 females and 1 male with Lambert 
N313400 E358900, 280m., VII.28.1991, 91- 
SRNP-1636; 2 females with Lambert 
N313400 E358900, 280m., VI.18.1994, 94- 
SRNP-4497, ex. Mauduca baruesi (paraty- 
pes deposited in USNM, BMNH, and 
CNC). 

Etyuiologi/. — This species is named in 
honor of Walter Bonilla Vasquez in special 
recognition of his dedicated management 
of the Accounting Office for the Area de 
Conservacion Guanacaste. 



226 



Journal or Hmvienoptera Research 




Figs. 88-89. Eiiplcctnis scanning electron micrographs. 88, E. xioiiinnw, Male scapes. 89, £. zvaltcri, Hind tibia 
and tarsi. 



Etiplectrus xiomarac Schauff, new 
species 

(Figs. 8, 9, 11, 19, 36, 85-87) 

Diagnosis. — Face below toruli dark 
brown (as in Fig. 13); mandibles yelk^w; 
hind coxa dark brown (Fig. 11) and hind 
femur with distal half brown; ss2 setae ab- 
sent between posterior ocelli (Fig. 83); ma- 
lar suture absent below eye. Male antenna 
with scape white, slightly swollen (Fig. 36, 
88), with narrow, elongate sensory area 
containing 2 irregular rows of sensillae ex- 
tending about 1/3 length. 

This is one of the small group of species 
with a dark face and dark hind coxae. It 
is most similar to £. janiici which, like 
xiomarac, lacks small setae between the 
posterior ocelli. Euplcctnis xiomarac can be 
separated by the presence of yellow man- 
dibles (mandibles brown in jamici) and 
lack of a malar suture (malar suture pre- 
sent in jamici). 

Descriptio}!. — Female. Body length 1.8- 
2.2 mm. Color: body mostly black except 
the following: face below and between to- 
ruli brown; antenna with scape yellow, 
flagellum yellow or light brown; mandi- 
bles yellow; legs yellow-brown except 
hind coxa mostly dark brown, hind femur 
with distal half brown; dorsal metasoma 
with large central yellow area extending 
from just behind petiole posteriorly about 
1/2 length of dorsum, becoming darker 
brown posteriorly, lateral margin brown. 



ventral metasc^ma yellow in anterior half, 
then dark brown. Head. Dorsally with mi- 
nor seta ss2 absent between posterior ocel- 
li (rarely with 1) (Fig. 85), all setae Sl-6 
present, setal row sr2 present as 1-2 irreg- 
ular rows of 8-10 setae reaching the bot- 
toni of the eye; occipital carina weak; 
width of eye: width of face:38:12, posterior 
margin eye of not nearly contiguous with 
postericir margin of head over most of 
length. Ratio of MS:EH 18:28; lateral ocel- 
lus more than 1 diameter from eye (OD: 
OOD 6:10). Face below eyes rounded, not 
abruptly narrowing. Vertex under anterior 
ocellus reticulate. Toruli separated by 
about 2.1 X their own diameters. Malar su- 
ture absent below eye. Area under eye 
lightly reticulate. Scape 4X as long as 
wide. Ratio of funicular segments 12:11:11: 
11:17, width 6 at Fl to 7 at club, flagellar 
segments with small whorls of brown se- 
tae basally. Mcsosoma. Pronotum anterior 
to transverse carina with scattered setae, 
finely rugosely reticulate, posterior to ca- 
rina more openly reticulate and shiny. Me- 
soscutum (Fig. 86) reticulate, becoming 
more smooth, and shiny pc^steriorly. Mid- 
lobe with median carina fading over an- 
terior 1 /4, otherwise complete, not notice- 
ably sunken posteriorly, with no small se- 
tae antero-laterally, posterior setae even 
with surface. Dorsal axillar/scutellar mar- 
gin with broad, nearly straight deep fur- 
row with flat bottom. Axillae shiny, open- 



Volume 10, Number 2, 2001 



227 



ly reticulate. Scutellum shiny, lightly retic- 
ulate to alutaceous, smc^oth along poste- 
rior margin, pc^inted at anterior margin 
with axillae. Metanotum bordered anteri- 
orly by a narrow band of small alveoli, 
medially shiny and lightly reticulate (Fig. 
87). Propodeum laterad of median carina 
lightly reticulate, shiny to the step-like pli- 
ca, niedian carina with anterior cup-like 
flange nearly triangular and invaginated, 
with transverse carina. Area around spi- 
racle openly reticulate laterally, spiracle 
slightly raised above surface and slanted 
so that opening is at a slight angle to the 
surface, with antero-lateral flange large, 
obvious, with 9-12 setae laterad and be- 
low spiracle. Posterior margin of propo- 
deum without deep alveolus at posterior 
margin. Petiole in dorsal about as wide as 
long (12:13), rugose dorsally. Mctasomn. 
Ovate, about 1.2-1.5X as long as wide. 
Legs. Ratio of hind tibial spur l:spur 2: 
tarsus 1:T2:T3:T4. 35:26:21:15:10:16. Fore- 
wing. Hyaline, about 2.3 X as long as 
wide. Costal cell with 2 irregular rows of 
setae ventrally. Venatit^n yellow to white, 
ratio of postmarginal: stigmal (25:18). 

Male. — Siniilar to female except: bociy 
length 1.9 mm; antenna with scape white, 
slightly swollen (Fig. 19, 36, 88), with nar- 
row, elongate sensory area containing 2 ir- 
regular rows of sensillae extending abc^ut 
1/3 length; funicle ratios 11:11:13:13:17 
width 5-6, with scattered or whorled 
semierect brown setae on each flagellom- 
ere. 

Hosts. — Hemiceras clarki, H. coreiua, H. iii- 
grescens, Rosenm ntteiiiiatn (all Notodonti- 
dae). 

Notes. — One female specimen has been 
observed with a single ss2 seta instead of 
the usual two. This further confirms that 
there may be some variation in this char- 
acter although overall it is c^uite stable. 

Distribution. — Known only from the 
ACG. 

Types. — Holotype female: Costa Rica, 
Guanacaste Prov., Area de Conservacion 
Guanacaste, Lambert N318500 E359850, 



240m., VIII. 20, 1993, 93-SRNP-4388, D.H. 
Janzen & W. Hallwachs. ex. Hemiceras clar- 
ki (deposited in INBIO). Paratypes: 3 fe- 
males with the same data as holotype; oth- 
er specimens with some of the same in- 
formation are except: 3 males and 2 fe- 
males with Lambert N318500 E359850, 
240m., VIII. 22. 1992, 92-SRNP-4609; 1 
male and 2 females with Lambert N3 18500 
E359850, 240m., VIII. 16. 1993, 93-SRNP- 
4384; 3 females with Lambert N313500 
E359850, 240m., VIII. 22. 1992, 92-SRNP- 
4614; 1 female with Lambert N318500 
E359850, 240m., VIII.23.1992, 92-SRNP- 
4642; 1 female with Lambert N318500 
E359850, 240m., VIII.22.1992, 92-SRNP- 
4602; 3 females with Lambert N318500 
E359850, 240m., VIII. 23. 1992, 92-SRNP- 
4652, ex. Rosenia atteniiata (deposited in the 
USNM and BMNH, INBIO, and CNC). 

Etymology. — This species is named in 
honor of Xiomara Driggs Valerin in spe- 
cial recognition of her dedicated manage- 
ment of the Human Resources office of the 
Area de Conservacion Guanacaste. 

Eiiplectriis zamorai Schauff, new species 

(Fig. 22) 

Diagnosis. — Face below toruli dark 
brown or black (as in Fig. 15); manciibles 
brown; seta S4 and seta ss2 absent (as in 
Fig. 85); funicle yellow except last two seg- 
ments dark brown (Fig. 22); malar suture 
absent below eye; setal line sr2 reduced; 
anterior metanotum without alveoli; pro- 
podeum laterad of median carina smooth. 

The contrasting distal antennal seg- 
ments of this species readily set it apart 
from similar species such as arias and val- 
verdei which also have a dark face and 
dark hind coxae but unicolorous, or only 
gradually darkening, flagellar segments. 
Euplectrus furnius also has the apical fla- 
gellar segments contrasting with Fl, but 
that species has the ocellus more than 2X 
its diameter removed from the margin of 
the eye (just over IX in zamorai) and the 
face is about 4X as wide as the width of 
the eye (just over 2x in zamorai). 



228 



Journal, of Hymenoptera Research 



Description. — Female. Body length 1.7-2 
mm. Color: body mostly black except the 
following: antenna with scape yellow, fla- 
gellum yellow, last 2 flagellomeres brown 
or light brown; mandibles brown; legs yel- 
low except hind coxa brown; dorsal me- 
tasoma with large central yellow area ex- 
tending from just behind petiole posteri- 
orly about 2/3 length of dorsum, becom- 
ing darker brown posteriorly, lateral 
niargin brown, ventral metasoma yellow 
in anterior half, then dark brown. Head. 
Dorsally with minor seta ss2 absent be- 
tween posterior ocelli, seta S4 absent, setal 
row sr2 present but reduced to 1 short 
row of 3-4 setae with an additional seta 
or two near bottom of eye; occipital carina 
weak to absent; width of eye: width of 
face: 33:14, posterior n"iargin eye of not 
nearly contiguous with posterior margin 
of head over n"iost of length. Ratio of MS: 
EH 15:28; lateral ocellus more than 1 di- 
ameter from eye (OD:OOD 6:8). Face be- 
low eyes abruptly narrowing. Vertex un- 
der anterior ocellus reticulate. Toruli sep- 
arated by about 2x their own diameters. 
Malar suture absent below eye. Area un- 
der eye lightly reticulate to alutaceous. 
Scape 5x as long as wide. Ratio of funic- 
ular segments 12:13:13:12:16, width 5 at Fl 
to 6 at club, flagellar segments with small 
whorls of brown setae basally. Mcsosonia. 
Pronotum anterior to transverse carina 
with scattered setae, finely rugosely retic- 
ulate, posterior to carina more nearly 
smooth with slight open reticulation and 
shiny. Mesoscutum reticulate, becoming 
more smooth, and shiny posteriorly. Mid- 
lobe with median carina nearly complete, 
fading at extreme anterior margin, not no- 
ticeably sunken posteriorly, with no small 
setae antero-laterally, posterior setae even 
with surface. Dorsal axillar/scutellar mar- 
gin with narrow, nearly straight deep fur- 
row without flat bottom. Axillae nearly 
smooth, shiny, with faint open reticula- 
tion. Scutellum shiny, very lightly aluta- 
ceous, smooth along posterior margin. 
Metanotum anteriorly nearly contiguous 



with scutellum, without obvious narrow 
band of small alveoli, medially shiny and 
smooth. Propodeum laterad of median ca- 
rina shiny and smooth to the step-like pli- 
ca, median carina with anterior cup-like 
flange rounded and invaginated. Area 
around spiracle reticulate, spiracle slightly 
raised above surface and parallel to the 
surface, with antero-lateral flange present, 
with 6-8 setae laterad and below spiracle. 
Posterior margin of propodeum without 
deep alveolus at posterior margin. Petiole 
in dorsal as wide as long (12:12), rugose 
dorsally. Mctnsoiun. Ovate, about 1.1-1.5X 
as long as wide. Legs. Ratio of hind tibial 
spur l:spur 2:tarsus 1:T2:T3:T4. 31:22:22: 
11:8:14. Forewing. Infuscate light brown, 
about 2.3 X as long as wide. Costal cell 
with 1 irregular row of setae ventrally. Ve- 
nation yellow light brown, ratio of post- 
marginal: stigmal (22:13). 

Mnlc. — Unknown. 

Hosts. — Unknown. 

Distribution. — Known only from the 
type localities. 

Types. — Holotype female: Costa Rica, 
Heredia, Chilamate, 75m., XII. 1989, 
111.1989, Hanson & Godoy (deposited in 
USNM). Paratypes: 2 females with same 
data as holotype except V. 1989; 1 female 
1X-X.1989; 1 female Guanac, Est. Pitilla, 9 
km S Santa Cecilia, 700m., V.1988, P. Han- 
son (deposited in USNM). 

Eti/niologif. — This species is named in 
honor of Luis Federico Garita Zamora in 
recognition of his many years of steadfast 
and enlightened managment of field op- 
erations and construction managen^ent for 
the Area de Conservacion Guanacaste. 

ACKNOWLEDGMENTS 

We thtink Tami Cdilow for teclmical assistance. 
Paul Hanson, Univ. of Costa Rica, kindly allovvecl us 
access to his collections of specimens. Drs. G. Zolne- 
lowich, S. Lingafelter, D. Smith and M. Gates re- 
viewed the manuscript and provided many helpful 
comments. This study was supported by NSF grants 
BSR 90-24770, DEB 93-06296, DEB 97-05072, BSR 80- 
11358, BSR 83-07887, and BSR 87-06155. This study 
would not have been possible without the support of 



Volume 10, Number 2, 2001 



229 



a number of Lepidoptera systematists and the rearing 
support of the paraecologists cited at http://janzen. 
sas.upenn.edu. 

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Dangerfield, P. C, J. B. Whitfield, M. J. Sharkey, D. 
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Journal of Hymenoptera Research 



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sion of the tribe Euplectrini of Sri Lanka (Hy- 
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new genera and species of Eulplectrini (Hyme- 
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]. HYM. RES. 
Vol. 10(2), 2001, pp. 231-244 

The Southern African Wasp Genus Handlirschia Kohl, 1897 
(Hymenoptera: Apoidea, Sphecidae, Bembicinae) 

Michael Ohl 

Museum ftir Naturkunde der Humboldt-Universitat Berlin, Institut fiir Systematische Zoologie, 
Invalidenstr. 40, D-10099 Berlin, Germany, E-mail: michael.ohl@rz.hu-berlin.de 



Abstract. — The poorly known digger wasp genus Hniuilirschia is revised for the first time, and 
two species are recognized, ncthiops (Handlirsch, 1889) and scoliiicfoiiiiis (Arnold, 1929); the latter 
is newly transferred from Stizus. Handlirschia tricolor Gess, 1973, is synonymized with scoliaeformis 
and a lectotype is designated for Stizus scoliaeformis. The revision includes diagnoses, descriptions, 
illustrations of significant characters for each species, and a distribution map. The phylogenetic 
position of Hamilirschia within the Bembicinae is discussed. 



Handlirschia Kohl, 1897 is a poorly 
know^n genus of digger wasp from south- 
ern Africa that u^as described for the sin- 
gle species, Stizus ncthiops Handlirsch, 
1889. The latter was based on a single 
male from eastern South Africa that re- 
mains the only specimen known. Gess 
(1973), without having seen the holotype 
of acthiops, described a second species, 
Hamilirschia tricolor, based on five speci- 
mens, also from eastern South Africa. 

As the first step toward a comprehen- 
sive revision of Stizus, I have compiled the 
original descriptions of species assigned to 
that genus. Based on the published de- 
scription and figures, Stizus scoliaeformis 
Arnold, 1929, seemed to be unusual in 
lacking diagnostic characters of Stizini and 
Bembicini, such as an elongate submargin- 
al cell I (Bohart and Menke 1976, Ohl 
1999). Examination of the type series 
showed that Stizus scoliaeformis actually 
belongs in the genus Hamilirschia within 
Gorytini (sensu Bohart and Menke 1976), 
and that Stizus scoliaeformis and Hamilir- 
schia tricolor are synonyms. Since then, 
roughly 50 more specimens of Hamilir- 
schia, all scoliaeformis ( = tricolor), came to 
my attention. These findings prompted 
me to revise the genus in order to evaluate 



its species composition, provide diagnos- 
tic characters of the included species, and 
evaluate its phylogenetic relationships. 

Diagnosis sections are not provided 
here since only two species are involved, 
and their defining characters are present- 
ed in the key. 

TECHNICAL TERMS 

Most morphological terms follow Bo- 
hart and Menke (1976). However, a few 
are explained here for convenience. 1 fol- 
low Melo (1999) in adopting the terminol- 
ogy of Smith (1970) for male genitalia. 
Gonapopln/sis: penis valve of Bohart and Menke 

(1976).' 
Goiiocoxitc: gonostyle of Bohart and Menke 

(1976). 
Metapostnotunr. usually referred to as 'propo- 
deal triangle', 'triangular area' or 'propodeal 
enclosure' in Apoidea, but in fact the meta- 
thoracic poshiotum that is fused to the true 
propodeum (Brothers 1976). In Handlirschia 
the metapostnotum is more or less triangular 
and extends slightly onto the posterior sur- 
face of the propodeum. 
Placoids and tyloids: I follow Bohart and Menke 
(1976:23-24) in distinguishing two kinds of 
specialized regions on male antennae. Pla- 
coids are "platelike, flat, or curved areas . . . 
that are . . . depressed below level of sur- 
rounding integument", whereas a tyloid is 



232 



Journal of Hymhnoptera Research 




Fig. 1. Hivhiliischia scoliacfoniiis, male, habitus. Namibia, Okahandja. 



defined as a "linear welt or cariniform swell- 
ing." 
Toiiihis (plural: toruli): antennal socket of Bo- 
hart and Menke 1976, i.e., the socket on the 
frons of the face upon which the scape of the 
antenna is articulated (Fig. 2a). 

All illustrations (except for Fig. 1, which 
is a traditional ink drawing) were pre- 
pared on a personal computer using Ado- 
be' programs: a pencil drawing made with 
a camera lucida was digitized with a scan- 
ner as a bitmap-based illustration and 
then imported into the Adobe Streamline 
4.0 program. This software converts bit- 
niap-based illustrations into vector-based 
illustrations, which were then modified in 
the Adobe Illustrator 7.0 program to pre- 
pare the final illustrations. 

Locality names are arranged in alpha- 
betical order within each country, district, 
or province, respectively. Coordinates 
were taken from various sources, espe- 
cially the catalogue of southern African 
place names by Leistner and Morris (1976) 
and the GEOnct names server of the 
NIMA Geographic Names Database (http: 
// gnpswww.nima.mil/geonames/GNS/ 
index. cfm). All coordinates follow the con- 
vention used by the 'Times Atlas of the 



World' (i.e., 21.55S 
21°55'S 16°08'E). 



16.08E instead of 



ORIGIN OF MATERIAL 

Institutional or personal collections in 
which the material is deposited are abbre- 
viated in the text as follows (names of con- 
tact person are in parentheses): 
AMGS Albany Museum, Grahamstown, 

South Africa (Fred W. Gess). 
BMNH British Museum (Natural Histo- 
ry), London (Christine Taylor). 
CAS California Academy of Sciences, 

San Francisco, USA (Wojciech J. 

Pulawski). 
CSE Personal collection of Christian 

Schmid-Egger, Berlin, Germany. 
MS Personal collection of Maximilian 

Schwarz, Ansfelden, Austria. 
NHMW Naturhistorisches Museum, Wien, 

Austria (Stefan Schodl). 
OHL Personal collection of Michael 

Ohl, Berlin, Germany. 
SAM South African Museum, Cape 

Town, South Africa (Margie A. 

Cochrane). 
USNM Smithsonian Institution, National 

Museum of Natural History, 



Volume 10, Number 2, 2001 



233 



Washington, D.C., USA (Mau- 
reen J. Mello). 

Genus Hamilirschia Kohl, 1897 

Handlirsdiia Kohl, 1897:425. Type species: Sphe- 
cius aefliiops Handlirsch, 1889, by monotypy. 

Diagnosis. — Handlirsdiia is a member of 
Gorytini (Bohart and Menke 1976), an as- 
semblage of 39 genera (Bohart 2000) that 
apparently lacks any apomorphy (see 
phylogenetic discussion below). The most 
important diagnostic features, which tax- 
onomically place Hamilirschia in Gorytini 
are the combination of two midtibial spurs 
(Fig. 2e), a keel-like basomedian ridge on 
sternum I, and submarginal cell 1 not un- 
usually elongate (Fig. 2b). Within Goryti- 
ni, Handlirschia belongs to the branch of 
genera with an oblique scutal carina (Fig. 
2c). Among these, it can be recognized by 
the lack of both a sternaulus and an omau- 
lus, and by the presence of spiracular 
lobes in the male. Additionally, the inner 
eye margins are almost parallel (Fig. 2a), 
whereas the margins converge ventrally in 
some other gory tin genera. 

Description. — A redescription of Handlir- 
schia is provided because Bohart and Men- 
ke (1976) based their generic diagnosis 
and description on the single known spec- 
imen of acthiops. They treated as distinc- 
tive for Handlirschia some characters of 
male aethiops that do not occur in scoliac- 
formis (e.g., the prominent male sternal 
fimbriae and somewhat distorted apical 
flagellomeres). Included are those charac- 
ters that vary within other Gorytini but 
are constant within Handlirschia. Reference 
is made to Stizini and Bembicini, which 
also have spiracular lobes in the male. 

Head shape simple (Fig. 2a): eye inner 
margins almost parallel; frons with angu- 
lar, transverse swelling below ocelli, shal- 
lowly depressed above antennae; toruli 
well above frontoclypeal margin; suban- 
tennal sutures well developed, reaching 
frontoclypeal suture between anterior ten- 
torial pits; clypeus slightly, evenly convex. 



maximum width 2.5-2.8x median length; 
mandibles slightly curved subapically, 
with an inner preapical tooth; male anten- 
nae with tyloids and/or placoids. Pronotal 
collar (Fig. 2d) sloping gently, separated 
from scutum by transverse groove, topped 
by a sharp edge (less so in scoliacforniis); 
scutum with well-defined oblique scutal 
carina (Fig. 2c); omaulus, sternaulus, epi- 
sternal sulcus and acetabular carina ab- 
sent. Hindleg arolium smaller than other 
arolia (Fig. 2f,g); mid tibia with two prom- 
inent spurs (Fig. 2e). Wings (Fig. 2b) in- 
fumate; pterostigma ill-defined, parallel- 
sided, posterior margin straight; both re- 
current veins received by submarginal cell 
II; jugal lobe larger than tegula; hindwing 
media diverging before cu-a. Propodeum 
without spiracular groove; propodeal 
hindcorners projecting, hindface concave. 
Metapostnotum an ecquilateral triangle; 
median propodeal pit forming longitucii- 
nal groove that extends from posterior 
third of metapostnotum slightly onto pro- 
podeal hindface and that is delimited by 
distinctive carinae; the latter project onto 
the propodeal hindface almost down to 
the propodeal orifice. Tergum VII with 
large spiracular lobe (Fig. 5c). Sternum I 
with median carina distally (in addition to 
much larger basomedian carina) and 
many oblique to longitudinal rugulae; 
sternum II with basal hump (Fig. 1) and 
prominent, deep, transverse groove ante- 
rior to it; sternum VII largely reduced to 
a membranous sclerite between spiracular 
lobes (Fig. 5c). 

Unfortunately, the genitalia and meta- 
somal segment VIII of the holotype of ac- 
thiops were lost after Handlirsch (1889:469) 
studied them. He described them as fol- 
lows: "The genitalia are almost the same 
as in the preceding species [which is a 
Sphecius in the sense of Bohart and Menke 
1976], except for the sagitta lacking the 
outer point" (my translation). Segments 
VII-VIII and the genitalia were dissected 
and originally glued onto card bottom, 
which is pinned under the specimen. Only 



234 



Journal of H^menoptera Research 



segment VII (and most of the right anten- 
na) is still present. Only two generic char- 
acters can be extracted from Handlirsch's 
description and scolincfonuis. Obviously 
sternum VIII is indeed narrowed to a 
sharp spine (Figs. 1, 5d, e). Apparently, 
the "outer point of the sagitta" [ = vol- 
sella] is the cuspis, which also lacks in sco- 
Uacfoimis (Fig. 3a, b). Absence of a cuspis 
is thus a generic character of Hninilirschin. 
The alleged similarity to Sf)hccius is not in- 
formative, because there are remarkable 
differences in the male genitalia between 
Hamilirscliin scolincfoi'Diis and the species of 
Sp^htxiiis that I have studied {tvitcinuitus, 
griDidis, hognrdii, pectornlis, spcciosiis, and 
spectabilis). 

Phylogenetic Position. — A few pre-cladis- 
tic hypotheses on the relationships of Hiiii- 
dlirscliin have been published. Handlirsch 
(1889:467) placed nethiops in Spheciits, 
pointing out that "within that genus, ne- 
thiops represented a group of its own" (my 
translation). Kohl (1897) established a 
nionotypic genus, Handlirschin for actiiiops, 
which he supposed to be "intermediate 
between Sphecius and Stiziis" . He gave a 
list of characters to differentiate Hmidlir- 
scJiin from the latter two genera and con- 
cluded that "apparently, Handlirschia is 
closer to Stiziis than to Spiiecius" (Kohl's 
Stiziis included Beinhcciiuis, to which he 
probably referred). Arnold (1929) placed 
scoliaeformis in Stiziis and commented: 
"Quite unlike any other species of the ge- 
nus and perhaps deserving to rank as the 
type of a subgenus" (Arnold 1929:318). 
Gess (1973) compared his new species, tri- 
color, only with Sphecius, thus implying 
placement in the Gorytini. Bohart and 
Menke (1976) emphasized the similarity of 
Handlirsciiia to Sphecius and Kohlia and 
considered Hmuilirschia as the most basal 
branch of those Gory tine that have an 
oblique scutal carina (Bohart and Menke 
1976:509, Fig. 155). Furthermore, they 
"put it on a separate line of evolutionary 
development in the general direction of 
the Bembicini and Stizini ... In fact Him- 



dlirschin would probably be put in Stizini 
except for the rather typical gorytin wing 
venation" (Bohart and Menke 1976:509). 

A comprehensive cladistic evaluation of 
the phylogenetic position of Haudlirschiti is 
beyond the scope of the present paper. 
Handlirschin is a member of Gorytini (sen- 
su Bohart and Menke 1976, Bohart 2000) 
within the clade Bembicinae (sensu Melo 
1999). As Bohart and Menke (1976) indi- 
cated, the Gorytini are a group of genera 
united only by symplesiomorphic fea- 
tures. Therefore, the tribe is probably a 
paraphyletic assemblage (Fig. 3a). Alex- 
ander's (1992) phylogenetic analysis of the 
Apoidea did not evaluate the phylogenetic 
status of the Gorytini, because he used 
tribes as terminal taxa. Melo (1999) ana- 
lyzed the relationships within Apoidea on 
the level of genera representing most of 
Bohart and Menke's tribes. Although he 
used only two representatives of the re- 
markably heterogeneous Gorytini {Hopli- 
soides and Ochleropitern), his results support 
the assumption of its paraphyly. 

Nemkov and Lelej (1996) presented a 
cladistic analysis of Gorytini (Fig. 3b). 
They did not include any outgroup (e.g., 
Stizini, Bembicini) in their study, howev- 
er, so that they could not test the phylo- 
genetic status of Gorytini. They studied no 
specimens of Hniuilirschia and relied on 
characters listed by Bohart and Menke 
(1976), some of which are lacking in sco- 
liaeformis. They established a new subtribe, 
Handlirschiina. This subtribe is not rec- 
ognized here, because it is a redundant 
name for an already named clade, Hau- 
dlirschia. Recognition of Handlirschiina 
would not be an improvement neither for 
taxonomic nor for phylogenetic purposes. 
Nemkov and Lelej placed Hniidlirsciiin as 
the basal branch of the assemblage of gor- 
ytine genera with an oblique scutal carina 
(Fig. 3b). This is not surprising because an 
oblique scutal carina is indeed a charac- 
teristic of most Gorytini (except for five 
genera). However, Nemkov and Lelej did 
not consider the potentially significant fact 



Volume 10, Number 2, 2001 



235 




Fig. 2. Handlirschin. Some generic characters (drawn from Namibian specimens of scoliacfaniiis). a. Fore and 
hindwing. b. Head frontally. c. Oblique scutal carina (osc), left posterior edge of scutum (sc) (tg = tegula). d. 
Anterior part of tfiorax (arrows indicate gently sloping pronotum and sharp-edged posterior groo\'e, respec- 
tively), e, Apex of left midtibia with two midtibial spurs, i, Hindtarsomeres IV-V. g, Foretarsomeres IV-V 
(female). 



236 



Journal of Hymenoptera Research 



Bembicini 



Stizini 
/ \ 

Bembecinus Stizus Stizoides 



/ 

remaining 
Bembicini 



\ 
Micro- 
Bicyrtes bembex 



remaining Gorytini with 
oblique scuta! carina 




remaining 
Bembicinae 



remaining Gorytini 
with oblique 
scutal carina 



Stizini 



Bembicini 



/^ 



\ 



/ 



"X 



IVIicro- remaining 

Bembecinus Stizus Stizoides Bicyrtes bembex Bembicini 



Bembicinae 
without oblique 
scutal carina 




remaining Bembicinae 
with oblique scutal carina 



VoLUMi^ 10, Number 1, 2001 



237 



that an oblique scutal carina is also pre- 
sent in Nyssonini, Stizini, and Bembicini 
within Bembicinae. 

The only unique apomorphy of Hninllir- 
sdiin that Nemkov & Lelej (1996) identi- 
fied is the presence of spiracular lobes of 
tergum VII (Figs. 4a, 5c). Besides Hainilir- 
schiii, this character occurs only in Stizini 
and most Bembicini (lobes missing in Bi- 
cyrtcs and Mkwbcmhcx), which together 
form a monophyletic group based on the 
presence of an unusually elongate sub- 
marginal cell 1. Since Nemkov and Lelej 
(1996) ciid not include Stizini anci Bembi- 
cini in their analysis, not surprisingly they 
considered the presence of a spiracular 
lobe as an apomorphy of Hniidlirschia. A 
preliminary alternative interpretation is 
depicted in Fig. 3c and explained below. 

The other three characters mentioned by 
Nemkov & Lelej (1996) as homoplastic 
apomorphies for Handlirsdiia are of less or 
no phylogenetic significance: (a) labrum 
prominent (weakly contrasting with the 
alternative character state "labrum incon- 
spicuous", and very likely modified sec- 
ondarily); (b) omaulus absent (omaulus 
missing in many other gorytine genera); 
(c) terga III, IV, and sometimes V with 
dense, apical fimbriae (present only in 
HaihilirscJiln acthiop^s). 

In summary. Kohl (1897) and more ex- 
plicitly Bohart and Menke (1976) were the 
first to imply a close relationship of Haii- 
dlirschin with Stizini and Bembicini, al- 
though they still placed the genus in Gor- 



ytini. As discussed above there is indeed 
a good reason to suppose that Hniidlirschia 
and Stizini + Bembicini together form a 
n"ionophyletic group, at least based on the 
striking presence of spiracular lobes. Fig. 
3c illustrates this preliminary hypothesis 
based on the assumption of a unique de- 
velopment of spiracular lobes, thus plac- 
ing Hniidlirschia as the sister group of Sti- 
zini + Benibicini. This tree agrees with a 
unique evolution of the elongate submar- 
ginal cell I in Stizini + Bembicini and the 
unique loss of the spiracular lobes in Bi- 
ci/rtcs and Microbciiibcx, but still implies a 
parallel evolution of the concave propo- 
deal hindface in Haudlirschia, Bciiibccinus, 
and Bicyrtes, respectively. Based on the six 
characters considered here, the hypothesis 
in Fig. 3c is more parsimonious than Bo- 
hart and Menke's hypothesis (Fig. 3a): 
there is no length difference for characters 
1-2 and 4-6 on the two trees, but spirac- 
ular lobes (character 3) were evolved only 
once in Fig. 3c rather than twice in Fig. 3a. 
However, this conflicting character polar- 
ization can only be more definitively re- 
solved in the frame of a cladistic analysis 
of the entire Bembicinae, taking into ac- 
count n^uch more characters and taxa. 

Life History. — Unknown. The presence 
of a female foretarsal rake suggests 
ground nesting. Flower records are known 
only for scoliacfonnis and are derived ex- 
clusively from the fieldwork of S.K. Gess 
and F.W. Gess, Grahamstown, South Af- 
rica. These data are given in the Life His- 
tory section of scoliacfonnis. 



KEY TO SPECIES OF HANDLIRSCHIA 
(The female of aethiops is unknown.) 

1. Metapostnotum srriooth, shiny, contrasting with coarsely, irregularly punctatorugose pro- 
podeal dorsum. Metapleuron and anterior part of propodeal side smooth, shiny. Black, 



Fig. 3. Hamilirschia. Three hypotheses on the phylogenetic relationships within the Bembicinae. a, Combined 
and redrawn after Bohart and Menke (1976: Figs 155, 181). b. Redrawn after Nemkov and Lelej (1996: Fig. 2). 
c. Preliminary hypothesis of the present paper. For discussion see Phylogenetic Position section. Character 
numbers: 1 = oblique scutal carina present, 2 = concave hindface of propodeum, 3 = presence of spiracular 
lobes, 4 = ocelli deformed, 5 = spiracular lobes reduced, 6 = elongate submarginal cell 1. 



238 



Journal of Hymenoptera Research 



appendages and face partly yellow-orange (males only?). Metasoma covered by distinctive 
erect setae that are two midocellar diameters long, pale on terga I-II, black on III-IV. Males: 
sterna III-V with short, erect, apical fimbriae (Fig. 4f); flagellomeres IV-XI shallowly exca- 
vated ventrally, with broad placoids; femora and tibiae conspicuously modified (see De- 
scription for details) aethiops (Handlirsch) 

Metapostnotum and propodeal dorsum coarsely punctatorugose to finely rugose (except 
for small impunctate area at apex of metaposhiotum). Metapleuron and anterior part of 
propodeal side microsculptured, dull. Thoracic dorsum and at least basal terga largely 
yellow-orange. Metasoma with short, appressed, golden setae less than one diameter long. 
Males: Sterna without fimbriae; flagellomeres V-IX at most slightly convex ventrally, with 
narrow, linear tyloids; legs unmodified scoliaeformis (Arnold) 



Handlirschia nethiops (Handlirsch, 1889) 

(Fig. 4) 

Spliecius Aetluops Handlirsch, 1889:467, male, in- 
correct original capitalization. Holotype: 
male. South Africa: "Caffraria" [eastern 
South Africa, see note below]: no specific lo- 
cality (NHMW), examined. — As Hiviiilirschia 
actliiops: Kohl, 1897:425 (new combination); 
Arnold, 1929:259 (Handlirsch's description 
translated into English); Bohart and Menke, 
1976:509 (listed); DoUfuss, 1989:9 (holotype 
in NHMW). 

Description. — (Based on holotype, a 
male.) Length 13.8 mm. Black with the fol- 
lowing yellow-orange: antennae, labrum, 
clypeus, frons below upper rim of toruli 
(somewhat extending above toruli at inner 
eye margin), and a narrow streak behind 
each eye. Flagellomeres II-XI somewhat 
distorted: II with oblique, inner ventral 
depression; III ventrolaterally with slight- 
ly convex, elongate, black spot, otherwise 
unmodified; IV with similar but much 
smaller spot and comma-shaped tyloid on 
ventral surface; V-VIII with depressed, 
shiny, ventral placoids; IX with a tiny, ba- 
soventral tyloici. Frons punctatorugose; 
vertex microsculptured, with scattered 
punctures, markedly punctured between 
ocelli and at posterior margin of vertex. 
Scutum shiny, with punctures about one 
diameter apart; punctures on scutellum 
and metanotum less than one diameter 
apart to subcontiguous. Mesopleuron 
coarsely and densely punctate, largely 
covered with long, dense, pale setae, al- 



most obscuring sculpture. Metapleuron 
and metapostnotum impunctate, shiny. 
Propodeal side impunctate and shiny be- 
fore spiracle, coarsely and sparsely punc- 
tate behind spiracle, many punctures 
more than one diameter apart. Propodeal 
hindface coarsely punctatorugose lateral- 
ly, with irregular, longitudinal carinae me- 
dially. Foretemoral venter markedly con- 
vex, with many erect, pale setae; foretibia 
with conspicuous, toothlike projection an- 
terobasally (Fig. 4e). Midfemur clublike 
(Fig. 4b), widest in apical half, depressed 
in basal half. Midtibia (Fig. 4b, c) deeply 
emarginate at spur insertion and pointed 
apically, with two prominent, apical 
spurs, inner surface with a prominent 
toothlike projection. Hindfemur parallel- 
sided, almost cylindrical (Fig. 4d). Setae 
pale on terga I-II, brown on terga III to 
VII. Tergal punctation coarse, sparsest on 
tergum I (many punctures 2-3 diameters 
apart), becon^iing denser from tergum II to 
VII. Setae pale on sterna I-III, brownish on 
IV-VII. Sternum II with rounded, basal 
hump. Sterna coarsely punctate to punc- 
tatorugose throughout. Sterna III-V with 
apical fimbriae (Fig. 4f), shortest on ster- 
num III (The setae of the sternal fimbriae 
are fused into compact triangles of re- 
markably similar size (Fig. 4f). This is 
probably a preservation artifact, and it is 
most likely the fimbriae originally formed 
compact, homogeneous rows). Sternum 
VII (Fig. 4a) small, sclerotized median part 
triangular, membranous laterally. Spirac- 



Volume 10, Number 2, 2001 



239 





f 



Fig. 4. Ha>tdlirfichia ncthiopis. a, Segment VII in oblique ventral view. b-c. Right foreleg, b. Anterior view, c, 
Oblique lateral view, d. Right hindleg, posterior view, e. Right midleg, posterior view, f, Sterna III-V, posterior 
margins with apical fimbriae. 



240 



Journal of Hymenoptera Research 














Fig. 5. I Iividllrschin scoliaefoiiiiis, male genitalia and metasomal segments VII-VIII. a. Genitalia, lateral view, 
b. Genitalia, ventral view (slightlv compressed to spread gonapophysis). c. Segment VII, oblique ventral view. 



Volume 10, Number 2, 2001 



241 



ular lobes of tergum VII long (Fig. 4a), ex- 
tending over more than half of total tergal 
length, approaching midline basally. Me- 
tasomal segment VIII and genitalia miss- 
ing. 

Type Localiti/ nini Collector. — Handlirsch 
(1889:469) gave the following information 
on the holotype of nethiops: "Siid-Afrika, 
Caffraria, Mus. Vindob. Coll. Winthem" 
(Mus. Vindob. = NHMW). Webster's Geo- 
graphical Dictionary defines Caffraria (or 
Kaffraria) as "a region of Eastern Cape 
Province . . . from Great Kei River on 
South to KwaZulu-Natal Province on 
North between the Drakensberg and the 
coast; largely equivalent to the main por- 
tion of former Transkei . . .". In the early 
19"' century, however, Caffraria was 
sometimes used in a broader sense, prob- 
ably covering most of eastern South Africa 
(F. Gess, pers. comm., Oct 2000). The ho- 
lotype is undated, but "Coll. Winthem" 
probably refers to the collection of Wil- 
helm von Winthem (1799-1847), whose 
Hymenoptera and Diptera material was 
transferred to the NHMW in 1852 (Horn 
et al. 1990). Von Winthem apparently nev- 
er traveled outside of Europe, but he com- 
municated with "close to 200 scientific 
correspondents" (Steetz 1848), who iden- 
tified, exchanged, and donated material, 
and he also purchased collections. Thus, 
the holotype of aetluops was obviously col- 
lected before 1847, but most likely not by 
von Winthem. 

Variation and Life History. — Unknown. 

Geographic Distribution. — Eastern South 
Africa is known. 

Material Examined. —SOUTH AFRICA: 
"Caffraria" ( = eastern South Africa): no 
specific locality (holotype male, NHMW). 



Haniilirschia scoliaeforinis (Arnold, 
1929), new combination 

(Figs. 1-2, 5-7) 

Stizus scoliacforDiis Arnold, 1929:317, female, 
male. Lectotype: male, Namibia: Kaokoveld, 
Warmbad (SAM), present designation (here 
designated in order to ensure the name's 
proper and consistent application), exam- 
ined.— Bohart and Menke, 1976:527 (listed). 

Haiidlirscliia tricolor Gess, 1973:103, female, 
male. Holotype: male. South Africa: Trans- 
vaal: Gravelotte, Beacon Ranch (AMGS), not 
examined. New synonym. — Bohart and 
Menke, 1976:509 (listed). 

Description. — Handlirschia scoliaeforinis 
was described in length by Gess (1973, as 
tricolor), whose paper should be consulted 
for more details. For coloration see Vari- 
ation section below. Frons microsculptu- 
red, impunctate, dull. Pronotal collar mi- 
crosculptured, with a few scattered ma- 
cropunctures. Mesopleuron coarsely, 
densely punctate medially. Metapleuron 
and propodeal side before propodeal spi- 
racle impunctate, dull; posterior half of 
propodeal side with coarse, scattered 
punctures that are denser posteriorly. Me- 
tapostnotum and hindcorners and dorsum 
of propodeum coarsely punctatorugose. 
Hindface of propodeum with two longi- 
tuciinal carinae. Midtibial spurs promi- 
nent, straight (Fig. 2e). Anterior hump of 
sternum II impunctate, microsculptured 
along midline. Sterna densely punctate, 
except sternum II obliquely punctatoru- 
gose laterally. 

Female. — Length 8.8-15.8 mm. Foreleg 
with foretarsal rake. Scutum, scutellum, 
and metanotum indistinctly punctatoru- 
gose. Foreleg arolium and tarsomere V 
markedly enlarged (Fig. 2g). Tergum I 



d. Segment VIII in ventral view, e. Apical spine of sternum VIII, lateral view. Abbreviations: dg = digitus; 
ga = gonapophysis; gc = gonocoxite; vo = volsella; SVII/SVIII = sternum Vll and VIII; TVII/TVIII = tergum 
VII and VIII. 



242 



Journal of Hymenoptera Research 



densely punctate throughout, punctures 
denser and shallower on following terga. 

M/7/('.— Length 10.0-16.9 mm. Flagel- 
lomeres III-VIl with linear tyloids, VllI 
with a circular tyloid, IX in most speci- 
mens without modifications, in some 
specimens with tiny, polished, basal spot. 
Scutum indistinctly punctatorugose, sin- 
gle punctures discernible toward posterior 
margin. Hindfemoral dorsum convex. 
Punctation of tergum I coarse, many punc- 
tures about one diameter apart, less than 
that on tergum II, shallower and denser 
on terga III to VI, tergum VII coarsely 
punctate. Spiracular lobes of tergum VII 
short, rounded, not extending beyond bas- 
al third of tergal length (Fig. 5c). Sternum 
VII membranous, bilobed apically (Fig. 
5c). Sternum VIII with a slightly curved, 
sharp spine (Fig. 5d, e). Gonocoxite with 
folded, membranous tip (Fig. 5a, b). Gon- 
apophysis compressed laterally (Fig. 5b), 
rounded in lateral view (Fig. 5a). Lateral 
margin of volsella embedded in and hard- 
ly discernible from basoventral part of 
gonocoxites (Fig. 5b); cuspis missing; dig- 
itus markedly sclerotized, narrow, with 
minute teeth dorsally (Fig. 5b). 

Variatioi. — Haiidlirschia scoliacforuiis 
shows a remarkable geographic color var- 
iation: almost all specimens from Namibia 
are extensively marked with yellow or 
yellow-orange, with a strongly contrasting 
black propodeum (Fig. 6a). Moreover, all 
terga are almost completely yellow-or- 
ange, with only the tergal margins nar- 
rowly black and with the band of tergum 
1 broken into two large spots. In some 
specimens one ore more of the basal terga 
have additional black markings, but the 
apical terga always have complete yellow 
bands (Fig. 6a). In contrast, specimens 
from eastern South Africa, including the 
types of tricolor, have the propodeum 
largely yellow-orange (Fig. 6b) or at least 
with some yellow-orange markings (Fig. 
6c). Additionally, the basal terga have 
more yellow than the terminal ones, 
which are usually all black (at least terga 




3 




6 c 




Fig. 6. Htvhilirschia scoliacforjiiis. Geographic color 
variation (based on males, but females exhibit the 
same tendency). Colors used: white = yellow-orange, 
black = black, grey = dark reddish-brown, a, Na- 
mibia, 30 km W Okahandja. b, South Africa, Gravel- 
otte (paratype male of H. tricolor), c. South Africa, El- 
lisras. 



V-VI in females and V-VII in niales) (Fig. 
6b, c). The sanie holds for coloration of the 
head, which has more black in eastern 
(Fig. 6c) than in western specimens (Fig. 
6a). One female from Rundu (Namibia) is 
interniediate in having a predominantly 
yellow-orange propodeum, a partly black 
face, and largely black, terminal terga. In 
addition to color, specimens from western 
Namibia average larger (females 11.2-15.8 
mm, males 12.8-16.9 mm long) than spec- 
imens from eastern South Africa (females 
8.8-13.0 mm, males 10.0-14.1 mm long). 

Although I did not study the holotype 
of tricolor and some other material from 
Namibia, housed in the AMGS, Fred Gess 
(pers. comm.) confirmed that these speci- 
mens correspond exactly to the above ob- 
servations. 

Life History. — All information on the life 
history of scoliacforuiis is derived from the 
fieldwork of Sarah K. Gess and Fred W. 
Gess (Grahamstown, South Africa), in- 
cluding identification of prey and flowers. 



Volume 10, Number 2, 2001 



243 



20' 



25° — 



30° — 




propodeum with 
red markings 

• propodeum black 



20° 



25° 



30° 



Fig. 7. Hividlirschia scoliacfonnis. Collecting localities. 



One male from Namibia has been collect- 
ed as prey of the asilid fly Siiphrolaunjrn 
bipiiiictiitn Loew. Floral records for scoliac- 
foniiis are: Amaranthaceae: Hernibstncdtin 
odornta (Burch.). Euphorbiaceae: Chninac- 
syce ^Inndiiligcm (Pax) Koutnik. Mollugi- 
naceae: Lifiicuin argute-carinatuui Wawra 
and Peyr., and L. nn/osotis E. Walter. 

GcograpyJiic Distribution (Fig. 7). — Namib- 
ia and Northern Province of South Africa. 

Material Examined (FSG is used here as 
an abbreviation for F.W. and S.K. Gess). — 
NAMIBIA: Karibib District: 84 km W of 
Okahandja on road to Karibib (21.55S 
16.08E), 2 Apr 1997 (visiting deep pink 
flowers of Hcrinbstaedtia odorata (Burch.) 
T.Cooke, Amaranthaceae), FSG (3 males, 
AMGS). Kavango Gebied: 100 km SW 
Rundu (17.56S 19.46E), 25 Jan 1993 (1 fe- 
male, MS). Khorixas District: 44 km from 
Helmeringhausen on road to Spes Bona 



(25.48S 16.23E), 16 Mar 1997 (visitiiig white 
flowers of Liiiicum itn/osotis E. Walter, Mol- 
luginaceae), FSG (2 males, 1 female, 
AMGS), same data, but prey of male Sti- 
pJirolaiiiyra bipuuctata Loew (Diptera: Asili- 
dae) (1 male, AMGS), same locality, 17 Mar 
1997 (visiting white flowers of Limeum nnj- 
osotis E. Walter, Molluginaceae) (1 male, 
AMGS), same data (visiting flowers of CJia- 
ijiaesi/ce glaiididigcra (Pax) Koutnik, Euphor- 
biaceae) (1 male, 1 female, AMGS). Kaross 
(19.30S 14.20E), "[South African] Mus. Ex- 
ped.", Feb 1925 (paralectotype female, 
SAM). Maltahohe District: Nomtsas 
(24.25S 16.51E), 18 Mar 1997 (visiting white 
flowers of Liuieuui argute-cariimtiuu Wawra 
and Peyr., Molluginaceae), FSG (1 female, 
AMGS). Okahandja District: Leeu River, 9 
km W Okahandja (21.58S 16.50E), 13 Feb 
1996, W.J. Pulawski (1 female, CAS); 30 km 
W Okahandja (21.55.56S 16.31. 61E), 1500m, 



244 



Journal of Hymenoptera Research 



Malaise-trap, 2-5 Mar 1997, M.O. Niehuis 
(7 males, 11 females, OHL; 1 male, 1 fe- 
male, CSE). Opuwo District: Warmbad ( = 
Warmquelle, 19.10S 13.49E), Koakoveld 
(probably a misspelling of Kaokoveld), 
"[South African] Mus. Exped.", Feb 1925 
(types of scoUaeformis (paralectotypes here 
designated): lectotype male, 2 paralectoty- 
pe males, paralectotype female, SAM). 
Otjiwarongo District: 18 mi NE Kalkfeld 
(20.45S 16.16E), 22 Feb 1996, WJ. Pulawski 
(2 males, 6 females, CAS; 1 female, 
BMNH); 25 km NE Kalkfeld (20.41S 
16.18E), 27 Feb 1996, WJ. Pulawski (2 fe- 
males, CAS); 15-20 km NW Otjiwarongo, 
3 Mar 1990, W.J. Pulawski (2 males, 4 fe- 
males, CAS). Outjo District: 24 km S Ka- 
manjab, 5 Mar 1990, W.J. Pulawski (1 fe- 
male, CAS); 18 km by road C40 from road 
C38 (20.02S 15.55E), 29 Mar 1997 (flying 
low amongst grass), FSG (1 female, 
AMGS). Tsumeb District: 10 km SE Tsu- 
meb (19.13S 17.42E), 8 Mar 1990, W.J. Pu- 
lawski (5 males, CAS; 1 male, MS). SOUTH 
AFRICA: Northern Province: Ellisras 
(23.40S 27.44E), 24 Dec 1973, H.N. Empey 
(1 male, AMGS). Gravelotte (23.57S 
30.37E), Beacon Ranch, Jan 1966, D.J. Broth- 
ers (types of tricolor: holotype male, 2 par- 
atype males, paratype female (referred to 
as "allotype" in Gess, 1973, and labeled ac- 
cordingly), AMGS; paratype male, USNM). 

ACKNOWLEDGEMENTS 

I thank Oliver Niehuis for donating Malaise-trap 
residues from Namibia, with twenty Hainilirschia sco- 
Uaeformis. I also thank Maximilian Schwarz for send- 
ing Handlirschia from his personal collection. Fig. 1 
was kindly prepared by Atilim Aynacioglu. I sincere- 
ly thank Wojciech Pulawski for sending me his syn- 
onymical and bibliographical catalog of Sphecidae. 
He and Fred and Sarah Gess provided information 
on specimens of Hmuilirschia in their collections. In 
particular, 1 thank Fred and Sarah Gess for allowing 
me to use their unpublished data on the life history 
of Handlirschia scoUaeformis. Wojciech Pulawski care- 
tuily checked the manuscript for spelling and consis- 
tency. My trip to South Africa was financiallv sup- 
ported by the Deutsche Forschungsgemeinscliaft 
DF( i in the course of a project on the phyk^geny and 
evolution of Apoidea (Wi 599/7; F^ainer Willmann). 



LITERATURE CITED 

Arnold, G. 1929. The Sphegidae of South Africa. Part 
Xll. Annals of the Transvaal Museum 13:217-319. 

Bohart, R. M. 2000. A review of Gorytini in the Neo- 
tropical region (Hymenoptera: Sphecidae: Bem- 
bicinae). Co)itributions on Entomology, Internation- 
al 4:1-259. 

Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps 
of the World. A generic revision. University of 
California Press, Berkeley, Los Angeles, London. 
1 color plate, ix -f 695 pp. 

Dollfuss, H. 1989. Verzeichnis der Grabwespentypen 
am Naturhistorischen Museum in Wien (Hyme- 
noptera, Sphecidae). Katalo;^e der iiussenschaftlich- 
en Sammlungen des Naturhistorischen Museums in 
Wien. Entomologie 7 (4):l-26. 

Gess, F. W. 1973. A new species of Hamllirschia Kohl 
(Hymenoptera: Sphecidae), a very poorly known 
genus from South Africa. Annals of the Cape Pro- 
vincial Museums (Natural History) 9: 103-107. 

Handlirsch, A. 1889. Monographie der mit Nysson 
und Bembex verwandten Grabwespen. IV. Sit- 
znngsherichte der Kaiserlichen Akadeniie der Wissen- 
schaften. Matheniatisch-Naturwissoischaftliche Clas- 
se. Abtheilung I 98:440-517, pi. I-II. 

Horn, W., I. KaWe, G. Friese, and R. Gaedike. 1990. Col- 
lectiones Entomologicae. Akademie der Landwirt- 
schaftswissenschaften der DDR, Berlin. 573 pp. 

Kohl, F. F. 1897. Die Gattungen der Sphegiden. An- 
)ialen des k.k. Naturhistorischen Hofnniseums 11 
(1896):233-516, pi. V-XI. [Dating after Review by 
A. Handlirsch. 1897. Verb. Zool.-Bot. Ges. Wien 
47:195-196, and Menke and Bohart. 1979. Proc. 
Entomol. Soc. Wash. 81:111-124] 

Leistner, O. A. and J. W. Morris. 1976. Southern Af- 
rican Place Names. Aiuials of the Cape Provhicial 
Museums 12: 1-565. 

Melo, G. A. R. 1999. Phylogenetic relationships and 
classification of the major lineages of Apoidea 
(Hymenoptera), with emphasis on the crabronid 
wasps. Scientific Papers. Natural History Museum 
of the Unii'ersity of Kansas 14:1-55. 

Nemkov, P. G. and A. S. Lelej. 1996. Phylogenetic re- 
lationships and classification of the digger wasps 
tribe Gorytini (Hymenoptera: Sphecidae, Nys- 
soninae). Far Eastent Entomologist 37:1-14. 

OhJ, M. 1999. A revision of Stizoides Guerin-Meneville, 
1844: taxonomy, phylogenetic relationships, bioge- 
ography, and evolution (Hymenoptera: Apoidea: 
"Sphecidae"). Mitteilungen ans dem Museum fiir Na- 
turkunde in Berlin, Zoologische Reihe 75:63-169. 

Smith, F. L. 1970. Evolutionary morpht)logy of the 
external insect genitalia. 2. Hymenoptera. Ainuds 
of the Entomological Society of America 63:1-27. 

Steet/, j. 1848. Nekrolog: Wilhelm von Winthem. En- 
tomologische Zeitung, Stettin ^): 104- 198 (with a 
postscript by Germar about tin- possibility to 
purchase the collection, p. IMS). 



J, HYM. RES. 

Vol. 10(2), 2001, pp. 245-250 

Nesting Biology of Isodontia costipennis (Spinola) 
(Hymenoptera: Sphecidae) 

LOURDES A. SOARES, LORENZO R. S. ZaNETTE, HeLCIO R. PiMENTA', ALDO GONgALVES, 

AND ROGERIO P. MARTINS 

Laboratorio de Ecologia e Comportamento de Insetos -ICB, Universidade Federal de Minas 
Gerais, P. O. Box: 486, Belo Horizonte, MG, Brazil (RPM E-mail: wasp@mono.icb.ufmg.br) 



Abstract .—Dm-m-y two years (10/92-10/93 and 10/95-10/96) the nesting biology of Isodontia 
costipennis (Spinola) was studied using trap nests in four distinct areas in the Campus of the 
Universidade Federal de Minas Gerais (UFMG), Belo Horizonte, M.G., Brazil. Females built linear 
series of one to six brood cells, weakly separated by wads of grass stems and filled with plant 
material (grass stems and fibers). Cells were provisioned mostly with nymphs of Tettigoniidae 
and Gryllidae. Wasp's eggs were laid on the cephalothoracic junction of the prey. The number of 
prey per nest varied from 1 to 18. Sixty-one adult wasps emerged from the 41 occupied trap nests 
(69'!'o females, 31% males) and the sequence of sexes in the cells was variable. Thorax widths of 
females were significantly larger than males (respectively 2.63 ± 0.22 mm and 2.37 ± 0.20 mm). 
An ichneumonid wasp {Mcssatoponis sp.) was the only natural enemy found. 



Isoiioutin (Patton) is a cosmopolitan ge- 
nus of solitary non-fossorial wasp that in- 
cludes 60 species (Hanson ancl Gauld 
1995). Females of this genus build their 
nests in natural cavities using many plant 
materials such as grass stems, flower pap- 
pus, moss and bits of wood; in one species 
bits of soil and charcoal are also used 
(Evans and Eberhard 1970; Bohart and 
Menke 1976). A number of important 
studies using the trap-nest method have 
been done on some species of Isodontia 
(Medler 1965; Lin 1966; Krombein 1967). 
hi addition, Piel (1933) studied in detail 
the Oriental species Isodontia iiigelliis 
(Smith). However, information about the 
Neotropical Isodontia species is scarce (Bo- 
hart and Menke 1976), especially for Iso- 
dontia costipciuiis (Spinola). The earlier re- 
ports on this species referred only to 
glimpses of nest construction and provi- 
sioning (Richards 1937; Berland 1929; Lin 
1966). Here we present new biological in- 



'In niL'moriam. 



formation about /. costipennis, especially 
on nesting behavior. 

MATERIALS AND METHODS 

Study sites. — This study was done on the 
Campus of the Universidacie Federal de 
Minas Gerais (19°52'S, 43°58'W 830 m), 
Belo Horizonte-MG, Brazil. The trap nests 
were placed in two different successional 
sites. Between October 1992 and October 
1993 the nests were placed in the "Esta^ao 
Ecologica" (site I), a preserved area since 
1969, that contains two secondary growth 
forest fragments, more than 40 years old, 
one "cerrado" fragment, one marshy area 
anei one grassland small area. 

From October 1995 to October 1996 a 
non-preserved area, also in the UFMG's 
campus, the "Prefeitura" (site II), was 
used for the observations. This area has 
some remains of forest, grassland and cer- 
rado mixed with exotic and ornamental 
species. 

Sampling. — Data were collected using 
trap nests (n = 4800), made of pieces of 
bamboo canes {PJiifllostacln/s sp.) 84 to 180 



246 



Journal of Hymenoptera Research 



■ Females 
D Males 



dilDll 



CNOJCNCOCOCOOOCOCOCOCOCOCOCOCO 

(j)(j)a><j)0)CT)CT)CT)ci>a)a)a><7)CT)a5 

t-t-t-OOOOOOOOOt-t--.- 

Month/Year 

Fig. 1. Number of adult Isodoiitia costipcnnis emerg- 
ing from the 13 nests of site I. 



7 
q5 6 
f 5 
i 4 

i 3 

< 2 
1 




I 



j ■ Females 
in Males 



nn 



■<a- ID CD r~- 

o o o o 
MonthA'ear 

Fig. 2. Number of adult Isodoiitin costipciuiis emerg- 
ing from 28 nests of site II. 



mm long and with 76 to 241 mm of inter- 
nal diameter. The bamboo cylinders were 
transv^ersely cut to provide a removable 
cover that allowed observations on larval 
development (Fig. 3). One side was closed 
using clay, and a hole of approximately 
6.5 mm was left in the other side. 

The trap nests were attached horizon- 
tally to wood sticks and trees at one and 
two meters height, forming two plots of 
600 sampling points in each studied area, 
one in the forest fragment and one in the 
cerrado fragment. Each sampling point 
had two trap nests (at 1 and 2m high), and 
was 10 meters away from the others. Each 
plot covered an approximate area of 
5000m-. 

All the nests were inspected monthly, 
and the occupied nests were collected and 
replaced by empty ones. In the laboratory, 
the collected nests were placed in trans- 
parent glass tubes closed with gauze and 
daily observed. 

Emerged adults of /. costipoiiiis were 
sexed and their thorax width was mea- 
sured with a digital pachymeter. This 
measurement was taken as the maximum 
distance between the external margins of 
the tegulae. After that, they were pined 
and placed in the UFMG's Insect Ecology 
and Behavior Laboratory reference collec- 
tion. 

Analysis. — All the analyses were carried 
out with Statistica for Windows (version 
4.3). The differences between internal di- 



ameters of occupied and non-occupied 
nests and between thorax width of males 
and females were tested by Student's t- 
test (Sokal and Rohlf 1995). The correla- 
tions between the nests length and diam- 
eter with thorax width; cocoon length; di- 
ameter and number of cells per nest were 
tested by Pearson's Correlation test. Dif- 
ferences between the amount of occupied 
nests at 1 and 2 meters high and propor- 
tion of sexes were tested using Pearson's 
Chi square. 

RESULTS 

Isodoiitin costipieunis occupied 13 trap 
nests in site I (= 0.3% out of 4800), 10 were 
at Im high and 3 at 2m high. Twenty-eight 
nests were occupied in site II (= 0.6 % out 
of 4800), 18 at 1 m high and 10 at 2 m. 
There were no differences between the 
number of occupied and non-occupied 
nests at each height, both, within sites and 
between theni (p = 0.41). In both areas 
only one generation of Isodoiitin costipciuiis 
per year was observed. Twelve females 
and 9 males emerged from the nests of site 
I and 30 females and 10 males from the 
nests of site II (Figs. 1 and 2). No differ- 
ences were found between the number of 
males and females emerged from each site 
(p = 0.15). Nests containing both sexes 
were found and the sequence of sexes in 
the cells was variable (Table 1). 

The average internal diameter of the oc- 
cupied nests of site I was 12.86 ± 3.84 mm 



Volume 10, Number 2, 2001 



247 



Table 1. The sex sequences of adults of hoiloutia costip>en)iis reared from trap nests (M = male, F = female, 
= sex unknown). 





iber 


Diite of emergence 






Cells per nc^ 


■.t 






Nest miin 


1 


2 


;i 


4 


- 


(1 


1 




5/8/92 


M 


F 


F 








2 




5/20/92 


F 


M 


M 








3 




7/3/96 


F 


M 


M 








4 




8/5/96 


F 


F 


F 








5 




10/26/95 


F 


F 


F 








6 




7/29/93 


F 


F 


F 








7 




7/29/93 


F 


F 


F 








8 




10/26/95 


M 


F 


F 








9 




8/7/96 


F 


F 


M 








10 




7/2/96 


M 


F 


M 








11 




10/26/96 


F 


F 


* 


F 






12 




10/27/96 


M 


F 




F 






13 




8/7/96 


M 


M 


* 


F 






14 




5/29/96 


M 


M 




F 


M 




15 




5/29/96 


F 


F 




F 


F 


M 


Females 


per 


cell ("o) 


60 


73 


60 


100 


50 






(range: 9.4-24.1 mm) and 10.57 ± 1.37 mm 
(range: 8.1-13.1 mm) to the nests of site II. 
No differences were found between sites 
(p = 0.0007). Non occupied-nests from site 
II presented internal diameters (14.92 ± 
2.86 mm) larger than the occupied ones (p 
= 0.0000), however no differences were 
found at site I. 

The average size of feniales from site I 
was bigger than the males (2.52 ± 0.2 n^m; 
2.33 ± 0.19 mm; p = 0.04). No differences 
were found between sizes of adults 
emerged from nests of site II (2.62 ± 0.28 
mm; 2.46 ± 0.19 mm; p = 0.09). No cor- 
relation was found between the internal 
diameter of occupied nests and the adults 
thorax width (r = -0.05; p = 0.6). 

Nest coustnictiou. — The females of Iso- 
doitia costipcuuis start nest construction, 
building a plug of tightly coiled plant ma- 
terial. Normally, two parts can be distin- 
guished in this plug, the first one made of 
small coiled shoot leaves (average length 
of the plug and standard deviation = 
11.83 ± 6.99 mm; n = 10) and the second 
made of tightly coiled leaf hairs (12.74 ± 
5.35 mm; n = 10). However some nests (n 
= 5) did not exhibit one of these parts. 



Subsecjuently, cells are constructed with 
sparsely coiled leaf hairs and the divisions 
between cells are made of compact plugs 
of leaf hairs (less compact than the initial 
plugs). The length of the divisions ranged 
from 5.5 to 23.26 mm (n = 11); in 6 nests 
a single undivided cell was observed. The 
number of cells per nest varied from 1 to 
6; 48% of them presented 2 cells. No cor- 
relation was found between the number of 
cells per nest and their length (r = 0.28; p 
= 0.3). An empty cell near the entrance 
was founci in 4 nests. Nest closure is done 
with another plug of firmly coiled leaf 
hairs (average length and standard devi- 
ation = 19.03 ± 2.92 mm) followed by 
loosely coiled leaf bits and ended with 
grass stems or dry brushwood tufts that 
can protrude until 30 mm from the en- 
trance hole (Fig. 3). 

Nest provisioning. — The number of prey 
per nest ranged from 1 to 18 (n = 21), usu- 
ally nymphs of one of the following En- 
sifera: Agraecidae, Copiphoridae and Pha- 
neropteridae. In one nest an adult of Try- 
gonidiidae (Grylloidea) was found. Only 
in two occasions, nests with mixed prey 
were found (Trygonidiidae and Copiphor- 



248 



Journal of Hymenoptera Research 




Fig. 3. Trap nest occupied by Isodoiitin costipciiiiis (1. 
dry brushwood tuft; 2. closure plug; 3. cell with prey; 
4. cell division; 5. cell with empty cocoon; 6. initial 



idae; Copiphoridae and Phaneropteridae). 
Six unconsumed prey (nymphs) remained 
alive and paralyzed over 7 days. Prey 
were placed venter up and the wasp's 
eggs were laid on their ventral cephalo- 
thoracic junction. 

Larval behavior. — All the observed larvae 
(n = 15) were active, constantly twisting 
and nioving their heads up and also open- 
ing and closing their mandibles until they 
touched the prey, then starting to con- 
sume them by biting their niesothoracic 
venter. Generally the larvae left behind 
only legs and heads of adults or nymphs 
in the last instar. 

On three occasions, larvae were ob- 
served moving from one cell to another. 
Once a larva, after eating all the prey in 
its cell, moved to a contiguous cell where 
there was a cocoon already and ate the re- 
maining prey that was there before spin- 
ning its own cocoon. In three others nests 
two cocoons were found in the same cell. 

hnuiatiirc iiistars. — Egg to adult devel- 
opment lasts 48 to 55 days in laboratory 
uncontrolled conditions. The eggs hatched 
in 1 to 3 days, and larvae took 5 to 12 days 
to start spinning their cocoons. The adults 
took 26 to 48 days to emerge. The cocoons 
were 15 to 17 mm long and 5 to 6 mm 
wide and were always positioned with 
their larger part toward the nest entrance. 
They had two layers, an internal brownish 
chitinous one and an external whitish fil- 
amentous one. No correlation was found 
between the cocoon size (diameter) and 
nest size (r = 0.12, p = 0.13). Almost 25% 
of the imn^iatures did not attain the adult 
stage. 

hitcractioii with otiier insects. — Seven in- 
dividuals of one species of Ichneumonidae 
wasp {Messatoporus sp.) were found in 5 
nests of /. costipcunis, 2 from site II and 3 
from site 1. In two other nests from site II, 
1 unidentified adult coleopteran was 
found with remains of /. costipciiiiis' co- 
coons. Nests from both sites were found 
containing ants {Caiuponotus spp.), how- 
ever only in one nest from site I evidences 



Volume 10, Number 2, 2001 



249 



of ant attack were found. Two males of 
Mcgachilc (Psciidoceiitroii) ciirvipcs (Smith) 
emerged from the same nest that one fe- 
male of /. costiphvniis emerged. 

DISCUSSION 

Isoiioitia costipviinis seems to be a locally 
rare species considering that only 0.3% 
(site 1) and 0.6% (site II) of the trap nests 
were occupied. However, the differences 
between the width of occupied and non- 
occupied nests suggest that the females of 
/. costipciuiis may prefer nesting cavities of 
specific size. Hence, considering that most 
of the unoccupied nests (78%) had an in- 
ternal diameter bigger than the occupied 
nest average diameter, it may be assumed 
that nest width was a liniiting factor for 
female selection. Fifty-eight percent of the 
trap nests occupied by Isodoiitia iiicxicaiia 
had an internal diameter of 6.4 mm and 
thirty-nine percent had an internal diam- 
eter of 7.9 mm (Medler 1965). The same 
author suggests that 4.8 mm would be the 
minimal diameter of the possible occupied 
nests. Such preferences probably could be 
related to the major efforts required by the 
females on the nest construction (Ainslie 
1922; Evans 1959). The differences in size 
between males and females, found only in 
wasps emerged from nests of site I, may 
be related to the small number of occu- 
pied nests and therefore not be a pattern 
for this species. Further studies will be re- 
ejuired to elucidate that question. 

The architecture of the observed nests of 
/. costipciuiis is similar to those from other 
species in the genus. Like 7. iiigcUa and /. 
pclopoiformcs, I. costipcuuis occasionally 
builds nests that could be considered in- 
termediate between the unilarval multi- 
cellular nests (as in /. clcgniis), and the 
niultilarval unicellular ones (as in 7. auri- 
pcs). In those nests, cell divisions are not 
well defined and more than one larva can 
be found in some cells (Bohart and Menke 
1976). Like /. uigclln and 7. clcgniis, I. costi- 
pciuiis builds (sometimes) multicellular 
nests with well defined partitions and a 



single larva per cell (Piel 1933; Krombein 
1967). The variety of nest types and ma- 
terials used in nest construction presented 
by 7. costipciuiis was also noted for 7. iiigcUn 
by Piel (1933) who presumed that a rela- 
tion between the materials used and local 
availability of certain plants might exist. 
Richards (1937) describes a nest of 7. cos- 
tipciuiis in a large curled-up leaf, with cells 
not clearly divided and filled with plant 
wool. The empty cells that were found 
near the entrances of four nests probably 
are vestibular. This type of cell may have 
served, in one period of nest evolution, to 
discourage parasites and predators from 
penetrating the stored cells (Krombein 
1967). 

The utilization of Agraecidae, Copi- 
phoridae and Phaneropteridae nymphs as 
prey has already been described for Iso- 
doiitia (Berland 1929; Medler 1965; Lin 
1966). However the provisioning, even if 
casual, with adults of Trigonidiidae, is a 
new observation for 7. costipciuiis. The 
number of prey placed in each cell by 7. 
costipciuiis also did not differ from other 
Isodoutia species. Bohart and Menke (1976) 
and Rau (1935) suggested that the number 
of prey per larva probably varies accord- 
ing to the size of the former. The variety 
of prey types used by 7. costipciuiis, may 
be directly related with the local avail- 
ability of prey (see Engelhardt 1928 and 
Medler 1965 for to other Isodoutia spp.). 
The niass provisioning and occasional 
mixed prey nests observed in 7. costipcnnis 
are also commonly found in the genus 
(Piel 1933; Medler 1965). 

The relatively low number of parasites 
founded in the studied nests, is atypical of 
other Isodoutia species. Five families of 
Diptera and three of Hymenoptera are 
commonly found parasitizing nests of 7s- 
odoiitia (Bohart and Menke 1976). Medler 
(1965), founded that 15% of the trap nests 
used by 7. mexicana produced parasites 
Sarcophagidae and Phoridae. Piel (1933) 
mentioned that some Stylopidae parasitize 
7. aiiripcs, I. Iiarrisi, I. iiigclln and 7. costipcii- 



250 



Journal of Hymenoptera Research 



nis, and are frequently found in the ori- 
ental species /. iiigelhi, but rarely in the 
South American species. No stylopized 
nest was founel in the present study. How- 
ever, the small number of parasites found 
in the nests of /. costipciiiiis may be only a 
result of the small number of trap nests 
occupied. Mixed nests with Mcgiicliilc 
were also reported for /. nicxicnua (Medler 
1965). 

The record of a single generation a year 
is also uncommon for Isodontin (Bohart 
and Menke 1976; Rau 1935). However, 
considering that in this study all the oc- 
cupied nests were removed to the labora- 
tory the length of development may have 
been changed. The high number of dead 
immature may also be related with to nest 
transference from field to laboratory. Like 
other species of the genus, sex sequences 
in the nests of /. costippcnis were variable 
(Medler 1965), but unlike other Isodontin 
spp., nests containing both sexes had cells 
with males preceding the cells with fe- 
niales. 

The nesting behavior of Isodojifln costi- 
pciinis is thus very similar to that of other 
Isodontin spp. However some distinguish- 
able aspects such as the utilization of Tri- 
gonidiidae as prey, the existence of only 
one generation a year and the "male be- 
fore female" sex sequences in the nests, 
should be noted. Although /. costipcnuis 
seems to be a locally rare species, further 
studies are required to elucidate this sup- 
position. 

ACKNOWLEDGMENTS 

Servio Tulio Pires Amarante identified /. costipcii- 
nis, Alejo Mesa the Orthoptera, Fernando Amaral da 
Silveira identified Megnchile (Pseudoceutroii) curvipcs, 
and Alice Fumi Kumagai identified Mcssatoponis sp. 
Frederico Drumond Martins helped in the field. Mar- 



cus Ferreira drew the trap nest (Fig. 3). Except for H. 
R. Pimenta, authors have scholarships from CNPq 
and Capes. FAPEMIG provided financial support. 
This is a contribution of the Program in Ecology and 
Wildlife Management of the Federal University of 
Minas Gerais. 

LITERATURE CITED 

Ainslie, C. N. 1Q22. Note on the nesting habits of 
Chlorioii clcgniis. The Cniiadiivi Eutoiiiologist, 269- 
270. 

Beriand, L. ]'-)29. Notes sur les Hymenopteres de 
France. Bulletin dc la Socictc Eiitoiiiolo;^iquc dc 
France 63-67. 

Bohart, R. M. and A.S. Menke. 1976. Spliccid Wasps of 
the World, a generic revision. University of Cali- 
fornia Press, Berkeley, 695 pp. 

Engelhardt, G. P. 1928. An observation on the breed- 
ing habits of Chlorion harrisi in Texas (Flymenop- 
tera). Bnlletiiii of tlie Brooklyn Entomological Societif 
23: 269-271. 

Evans, H. E. 1959. Isodontia, the grass carrying wasp. 
Nature Magazine 52: 237-239. 

Evans, H. E. and M. J. W. Eberhard. 1970. The Wasps. 
University of Michigan, Ann Arbor, 265 pp. 

Hanson, P. E. and 1. D. Gauld. 1995. The Hymenoptera 
of Costa Rica. Oxford University Press, 899 pp. 

Krombein K V. 1967. Trap-nesting Wasps and Bees: Life 
tiistories, Nests and Associates. Washington, D.C. 
Smithsonian Press, 570 pp. 

Lin, C. S. 1966. Bionomics of Isodontia mexicana, with 
a rex'iew of generic ethology (Hymenoptera: 
Sphecidae:Sphecinae). The Wasmann journal of Bi- 
ologi/ 24(2); 239-247. 

Medler, J. T. 1965. Biology of Isodontia (Miirrai/ella) 
me.xicana in trap nests in Wisconsin (Hymenop- 
tera: Sphecidae). Ainials of the Entomological Soci- 
ety of America 58(2): 137-142. 

Piel, O. S. J. 1933. Recherches biologiques sur les Hy- 
menopteres du Bas Yang-Tse (Chine). Annates de 
la Societe Entomologique de France 102: 109-154. 

Rau, P. 1935. The grass-carrying wasp, Chlo- 
rion(Isodontia) harrisi Fernald. Bulletin of the Brook- 
lyn Entomological Society 30: 65-68. 

Richards, O. W. 1937. Results of the Oxford Univer- 
sity Expedition to British Guiana, 1929. Hyme- 
noptera, Sphecidae and Bembecidae. Transactions 
of the Royal Entomological Society 86: 101-118. 

Sokai, R. R. and F. J. Rohlf. 1995. Biometry. 3" ed., W. 
H. Freeman & Co., New York, 887 pp. 



[. HYM. RES. 
Vol. 10(2), 2001, pp. 251-260 

Ultramorphology and Histology of the Foregut and Midgut of 

Pachycondyla (= Neopottera) villosa (Fabricius) Larvae 

(Formicidae: Ponerinae) 

FlAvio Henrique Caetano and Fernando Jose Zara 

(FHC) Departamento de Biologia, Instituto de Biociencias — Universidade Estadual 

Paulista— UNESP, P.O Box 199, 13506-900 Rio Claro (SP), Brazil; E-mail: 

fcaetano@rc.unesp.br; (FJZ) Departamento de Biologia, Faculdade de Ciencias e Letras — 

Universidade Estadual Paulista— UNESP, P.O. Box 335, 19806-900 Assis (SP), Brazil; 

E-mail:fizara@rc. unesp.br. 



Abstract. — We studied the foregut and midgut of larvae of Padnicondyla (= Neoponera) villosa 
(F.) using histological methods and the scamiing electronic microscope. In this paper we discuss 
the muscle layers of these regions, the origin of peritrophic matrix, the digestive cells secretion 
process, and the term proventriculus used in larvae. 



The digestive tract of hymenopteran lar- 
vae consists of the pharynx, esophagus, 
proventriculus, ventriculus and ileum 
(Wheeler 1926, Nelson 1924, Wheeler and 
Wheeler 1976). When compared to the 
adult digestive tract, the major difference 
is that the foregut of the larva has no crop 
and in the hindgut there is no clear differ- 
ence betw^een ileum and rectum. The lar- 
val hindgut consists of a short and narrow 
tube which widens at the end. The Mal- 
pighi tubules open into the anterior por- 
tion of the ileum. 

The ventriculus occupies most of the 
larval body and represents the major por- 
tion of the gut both in terms of length and 
diameter. The peritrophic matrix, which is 
well developeci iii larvae, is practically ab- 
sent in adults, where it is found in few 
cases (Caetano 1988, Caetano et al. 1986/ 
1987, Caetano and Hoffmeister 1987, Cae- 
tano et al. 1994). 

The major objective of the present study 
is to describe some ultraniorphological 
and histological aspects of larvae foregut 
and midgut of the PncJii/coudi/ln villosa (F.) 



and to present some ontogeiietic consid- 
erations. 

MATERIALS AND METHODS 

For ultramorphology study, P. villosn 
larvae were dissected in physiological sa- 
line for insects in a Petri dish covereci with 
colored wax. The digestive tract was re- 
moved and fixed in Karnovsky fluid for 
24 hours, dehydrated in a ascendiiig al- 
cohol series (70 to 100%), subjected to two 
acetone 100% baths of 15 minutes each 
and then critical pointed dried (Balzers 
CPD 030). After dehydration the material 
was placed on aluminum supports at- 
tached with double-faced tape and sput- 
tercoated with gold (in sputtering Balzers 
SD 050). The digestive tract was examined 
with a Jeol PI 5 SEM and photographed on 
Neopan SS 120 film. 

For historesin preparation, P. villosn lar- 
vae were fixed directly in 4% paraformal- 
dehyde in phosphate buffer (0.1 M, pH 
7.4). The material was then dehydrated in 
70, 80, 90 and 95% ethanol solutions for 20 
minutes and transferred to resin solution 
(JB4 — Polaron Instruments /Bio Rad) for 



252 



Journal of Hvmenoptera Research 




Fij^s. 1-2. 1, General view of k)regut. E = esophagus; SN = stomogastric nerve; LM = longitudinal muscle; 
I'V = proventriculus; V = ventriculus. Scale bar = 40|jim. 2, Detail of esophagus showing relationships with 
stomogastric nerve (arrow). CM = circular muscle. Scale bar ^ l()|a.m. 



Volume 10, Number 2, 2001 



253 




Figs. 3-5. 3, Froventriculus detail where longitudinal muscle occurs externally. Scale bar = 20|xm. 4, General 
view of mesenteron (V = ventriculus). MT = Malpighian tubules. Scale bar = 200|xm. 5, Detail of ventriculus 
wall showing thin and external longitudinal muscle and under these fibers the stronger circular muscle. T = 
trachea. Scale bar = 30|jLm. 



254 



Journal of Hymenoptera Research 










%4\ 



a^ 



EP 



H 



V 



.w 






MV 



-e*v^ 9 



Volume 10, Number 2, 2001 



255 



72 hours at 4°C. Finally the material was 
transferred to molds filled with resin con- 
taining a catalyze and sealed with metal 
support for microtomy. The blocks were 
cut in Sorvall JB4/Bio Rad microtome. The 
sections were stained with hematoxylin- 
eosin and photographed with Zeiss pho- 
tomicroscope. 

RESULTS AND DISCUSSION 

UUrnniovphology. — The esophagus is a 
short and narrow tube that dilates near 
the ventriculus to form the proventricu- 
lus (Fig. 1). This region is covered with 
a circular muscle layer (Fig. 2) that dc^es 
not resemble that observed in the adults 
of other ants (Caetano 1990). In the 
adults, the external muscle is visibly stri- 
ated and consists of oblique fibers that 
intercross, bifurcate or anastomose. In 
the larvae there were no such striations 
or fiber separation. The image is that of 
a sheath covering the entire organ. The 
circular muscle of the esophagus stop 
abruptly at the anterior border of pro- 
ventriculus and resemble those de- 
scribed to Soleuopsis iuvictn Buren (Pe- 
tralia and Vinson 1980) and other ant lar- 
vae (Valentini 1951). 

The stomogastric nerve runs along the 
entire larval esophagus towards which it 
emits small branches that penetrate the 
muscle sheath (Figs. 1 and 2), as is also 
observed in adults. 

The proventriculus appears as a protu- 
sion form the ventriculus surface (Fig. 1). 
This protrusion resembles that described 
by Caetano (1988), Caetano et al. (1986/ 



1987) and Tomotake (1990) for adults of 
the subfamily Ponerinae and called but- 
ton. Eisner (1957) and recently Tomotake 
et al. (1995a,b) have shown that in adults 
the prov^entriculus may have external lon- 
gitudinal muscle fibers, as also observed 
for some species of the subfamily Poneri- 
nae (Toniotake 1996). The presence of 
these longitudinal fibers in adults suggests 
that they may have retained this character 
froni larvae because the proventriculus is 
the only foregut region that has superficial 
longitudinal muscle fibers in both adults 
and larvae. Structurally this arrangement 
does not differ from typical figures of lar- 
vae presented in morphology textbooks 
(Wheeler and Wheeler 1976). The most 
clearly visible elements are the outer lon- 
gitudinal muscle fibers (Fig. 3), which 
were also observed in this portion of the 
digestive tract of P. villosa adults by trans- 
mission electron microscopy Caetano 
(1991). 

The ventriculus is elliptical, very wide 
(Fig. 4) and covered with weakly devel- 
oped longitudinal muscles and connec- 
tive tissue. The fibers of the circular mus- 
cle are located below these layers (Fig. 5). 
The location of these muscles follows the 
pattern known for adults of these and 
other ant species in which a web of amc^r- 
phous connective tissue is observed (Cae- 
tano unpublished data) through which 
tracheal branches penetrate the ventricu- 
lus (Fig. 5). 

According to Lappano (1958), Petralia 
and Vinson (1980) the ventriculus is the 
largest organ of the larva and appears to 



Figs. 6-9. 6, Longitudinal section of esophagus, showing thick cuticle (C) covering lumen. FB = fat body; L 
= lumen; N = nucleus; EP = esophagus epithelium. Scale bar = 10|JLm. 7, Transverse section in esophagus 
showing thick cuticle, cubic epithelium of esophagus and sheath of circular muscle. Scale bar = 20fi,m. 8, 
Section through proventriculus showing different epithelia of region. (EPT) transition epithelium, (SV) sto- 
modeal valve epithelium and (VEP) ventricular epithelium. In this micrograph we can observe that the tran- 
sition epithelium is producing the peritrophic matrix (arrow). The Malpighian tubules and the fat body are 
close to this region. PM = peritrophic matrix. Scale bar = 20|xm. 9, Transverse section in proventriculus. MV 
= microvilli; SV = stomodeal valve; L = stomodeal vahe lumen; C = cuticle; CCL = stomodeal chamber 
lumen. Scale bar = 20[xm. 



256 



Journal of Hymenoptera Research 




Fig. 10- 
micruvil 



3. 10, Vrntriculus epithelium with digestive cells (DC); cells have large median nuclei (N) and long 
li (MV) with smooth secretion droplets (SD) between them. Scale bar = 20(jLm. II, Detail ot apex ot 



Volume 10, Number 1, 2001 



257 



serve as a niould around which the other 
abdominal organs are orientated. This 
suggestion is confirmed in P. villosn lar- 
vae. 

Histologi/. — The esophagus has a wide 
lumen, cuboidal epithelium covered with 
a thick cuticle and a circular muscle sheath 
covering it externally (Figs. 6 and 7). Dur- 
ing feeding, the food bolus may be trans- 
ported through the esophagus by means 
of peristalsis, as occurs in S. iiivicta (Pe- 
tralia and Vinson 1980). Except for the 
wide lumen and thick cuticle, this descrip- 
tion agrees from that of adults of the same 
species (Caetano 1988) anci of all other 
adult ants studied so far (Walker and 
Clower 1961, Caetano and Lage Filhc^ 
1982, Caetano 1984, 1988, 1990). In adults 
the epithelium is thin and the muscle 
sheath less so evident. 

The proventriculus consists of columnar 
epithelium with basal nuclei and well de- 
veloped ensheathing longitudinal mus- 
cles. This portion of the foregut has two 
chambers: one formed by the epithelial 
portion of the foregut (cardiac or stomo 
deal valve), and the other located between 
the stomodeal valve and the proventricu- 
lus wall, whose lumen communicates di- 
rectly with the lumen of the ventriculus 
(Fig. 8). 

Thus, the proventriculus of P. villosa lar- 
vae appears to be formed by a prolonga- 
tion of the epithelium of the ventriculus 
towards the foregut and its internal por- 
tion, the stomodeal valve proper, is 
formed by a projection of the foregut to- 
wards the ventriculus (Fig. 8). Thus, these 
portion problably reflect a mixed devel- 
opmental origin: the internal part arises 
from the ectoderm of the foregut and the 
outer part from the endoderm of the mid- 



gut. The presence of outer longitudinal 
muscle fibers in the so-called larval pro- 
ventriculus, as observed in the ventricu- 
lus, is clarified. On this basis, we believe 
that in larvae the structure known as pro- 
ventriculus should be called "stomodeal 
chamber" because of its position and it 
harbors the stomodeal valve, in contrast 
with the adult proventriculus that does 
not have the stomodeal chamber (Caetano 
1988). 

The lumen of this region is lined with a 
thick cuticle organized in a trabecular for- 
mation, which does not resemble the cu- 
ticular arrangement of adults. The adult 
proventricular lining is organized into 
four (or more) mobile lips that are usually 
covered with spiniform cuticular struc- 
tures (Eisner 1957, Caetano et al. 1991, 
1998, Tomotake 1996). 

The ectodermal origin of the stomodeal 
valve was indicated by the presence of cu- 
ticle that does not stain in histochemical 
processes for proteins, as shown in trans- 
verse sections (Fig. 9). 

The ventriculus is a yellow or dark col- 
oured region visible in live larvae. It is a 
blind sac posteriorly, similar to what is ob- 
served in several species of higher Hy- 
menoptera as Apns ineUifcrn L. (Nelson 
1924), Eciton biirdwUi Westw. (Lappano 
1958), S. invictn (Petralia and Vinson 1980), 
Ectatoinum edentatiiin Roger (Zara and Cae- 
tano 1998) and some species of the sub- 
family Formicinae, Dolichoderinae and 
Myrmicinae (Valentini 1951), although 
Athias-Henriot (1947) reports it to be open 
in Monoiuoriuin. 

The ventricular epithelium is colum- 
nar and could be identified by one type 
of digestive cell. This cell has distinct 
round or oval pycnotic nucleus located 



digestive cells with long microvilli and secretion (S) close to apical membrane. Scale bar = 10|xm. 12, Detail 
of secretion droplets (SD) releasing from digestive cells. Note absence of microvilli in secretion droplet. Scale 
bar = 10|xm. 13, General view of ventriculus epithelium of old larvae showing large space around nucleus 
and secretion in apex. GC = Regenerative cell. Scale bar = 20|jLm. 



258 



JOURNAL OF HyMENOPTERA RESEARCH 



between the median and basal portion of 
the cell (Fig. 10) similar to E. biircliclli 
and S. iuvicta (Lappano 1958, Petralia 
and Vinson 1980, respectively). The apex 
of these cells is rounded, conferring a 
club-like shape on them, and usually ap- 
pears to contain large secretion vacuoles. 
The border bears long microvilli (Figs. 10 
and 11). The secretion is released, like a 
"budding" process in the form of vesi- 
cles containing granules that stain differ- 
entially with H-E (Figs. 10, 11 and 12). 
This process resembles that observed in 
the adult ventriculum of Pacln/condyln 
strintn Smith, but without microvilli sur- 
rounding the vesicles (Caetano et al. 
1994). This kind of release secretion is 
different than merocrine secretion de- 
scribed in Cntngli/pliis boniln/ciiui (Roger) 
larvae (Valentin! 1951). These cells could 
be producing digestive enzymes; how- 
ever, the absorbative cells were not ob- 
served. 

The nests of regenerative cells common- 
ly present in the ventriculus of adults are 
not observed in larvae (Caetano 1984, 
1988, 1990, Caetano et al. 1986/1987), but 
they occur as isolated nuclei close to the 
basal lamina (Fig. 13). 

The peritrophic matrix is clearly visi- 
ble in ventricular lumen (Figs. 14 and 15) 
and an accumulation of "membranes" is 
noted in its distal portion. Some of the 
membranes enclose food remains and 
some of them are empty (Fig. 16). In this 
region, the remnants of the peritrophic 
membrane (meconium) are accumulated 
for later elimination during pupation as 
also observed in S. iiwictn (Petralia and 
Vinson 1980). The anterior region of the 
proventriculus, more precisely the tran- 
sition epithelium (Fig. 8, arrow), seems 
to be responsible for the secretion of the 
peritrophic matrix which expands soon 
after leaving the cardiac valve (Fig. 14) 
and reaches the epithelium of the ven- 
triculus. This kind of peritrophic matrix 
formation is similar to that recorded for 
£. cdcntatuiii larvae (Zara and Caetano 



1998). In contrast, Petralia and Vinson 
(1980) decribed a second type of peri- 
trophic matrix produced along the entire 
ventriculus of S. iiivictn, but this does not 
occur in P. villosn. 

In adults of some ponerine genera (e.g., 
Ectntiviiiiin and Ncopuvicrn), Caetano (1988) 
reported a similar origin of the peritroph- 
ic matrix, which at first may be confused 
with the stomodeal valve itself. In the 
adults, however, the peritrophic matrix 
penetrates the median region of the ven- 
triculus and opens, appearing to origi- 
nate from inside the stomodeal valve. In 
the larvae studied here the peritrophic 
matrix opens immediately upon leaving 
the stomodeal valve, which serves as a 
mold. 



CONCLUSIONS 

a) The outer muscles present in the fore- 
gut and midgut of P. villosn larvae differ 
from those of adults by the absence of 
clearly transverse striations. 

b) The proventriculus and the ventric- 
ulus of the larvae have a external longi- 
tudinal muscle sheath but only the adults 
ventriculus shows it. 

c) The larval proventriculus is morpho- 
logically simpler than in adults; the pres- 
ence of cuboidal epithelium with micro- 
villi indicate a secretory function. 

d) The proventriculus term used for 
adults does not seem to be appropriate for 
the larval structure described here, we 
suggest the stomodeal chamber. 

e) The "button" located close to the ven- 
triculus anterior region previously de- 
scribed by other authors and present in 
adults of the subfamily Ponerinae is sim- 
ilar to the upper portion of the stomodeal 
chamber. 

f) The histological characteristics of the 
stomodeal chamber of larvae lead us to 
believe that this portion may originate 
from the endoderm as the ventriculus. 



Volume 10, Numbi-r 2, 2001 



259 




PM 



M 



16 



,=>i 



.J'- 



Figs. 14-16. 14, General view of veniriculus proximal region witli stomodeal valve aiui peritropliic ni.itri\ 
being released. Scale bar = lOirm. 15, Close up of peritrophic matrix showing its layers. Scale bar = 10p.m. 
16, General view of posterior region of ventricujiis with "meconium" (M). Scale bar = SOfim. 



260 



Journal oh Hymenoptera Research 



ACNOWLEDGEMENTS 

This work was svipported by FAPESP (Fundaqao 
de Amparo a Pesquisa do Estado de Sao Paulo) — 
Proc. ^^8/00576-8. 

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Athias-Henriot, C. 1947. Recherches sur les larves de 
quelques fourmis d'Algerie. Bullcti)i Biokn^iqiic dc 
In France ct de la Bclgiquc 81: 247-272. 

Caetano, F. H. 1984. Morfologia comparada do trato 
digestivo de formigas da suhfamilia Myrmicinae 
(Hymenoptera, Formicidae). Pnpcis Aviilsos dc 
Zoologia 35: 257-303. 

Caetano, F. H. 1988. Anatomia, histologia e histo- 
qui'mica do sistema digestivo e excretor de op- 
erarias de formigas (Fiymenoptera, Formicidae). 
NatumUa. 13: 129-174. 

Caetano, F. H. 1990. Morphology of the digestix'e 
tract and associated excretory organs of ants. 
Aplicd Mi/mcrcologi/., 1: 119-137. 

Caetano, F. H. 1991. Ultraestrutura do pescocjo do 
proventriculo de Ncopoucra villosn (Formicidae, 
Ponerinae). Proceeding's of the XIII Coloquio Socic- 
dnde Bmsllcini de Microscopin Eletrdiiicn, Caxambu 
p. 167. 

Caetano, F. H. and A. L. Lage Filho. 1982. Anatomia 
e histologia do trato digestivo de formigas do ge- 
nero Odontomnchiis (Hvmenoptera, Ponerinae). 
Natiiralia. 7: 125-134. 

Caetano, F. H., M. I. Camargo-Mathias and W. L. Ov- 
eral. 1986/1987. Anatomia e histologia compar- 
ada do trato digesti\'0 dc Dinopoiiem gigaiitea e 
Parapoiiera clnvntn (Formicidae, Ponerinae). Na- 
tumUa. 11/12: 123-134. 

Caetano, F. H. and R. M. Hoffmeister. 1987. Preseni^a 
de Membrana Peritrofica em Cniiipoiiotus riifipes 
(Hymenoptera, Formicidae). Proceedings oftlie. XI 
Coloquio Socicdade Brnsileira dc Microscopia Eletron- 
ica, Caxambu p.91-92. 

Caetano, F. H., D. Beig and J. D. Majer. 1991. Descri- 
qao do proventriculo de Mi/iucrcia sp (Formici- 
dae, Myrmicinae) ao microscopio eletronico de 
varredura. Proceedings of the XIII Coloquio Socic- 
dade Brasileira de Microscopia Eletronica, Caxambu, 
p. 157-158. 

Caetano, F. H., A. H. Torres, M. 1. Camargo-Mathias 
and M. E M. Tomotake. 1994. Apocrine secretion 
in ant, Pachycondyla striata, ventriculus (Formici- 
dae: Ponerinae). Ci/tobios 80: 235-242. 

Caetano, F. H., X. Espadaler and F. J. Zara. 1998. 
Comparative ultramorphology of the proventric- 



ulus bulb in two species of Mutillidae (Hyme- 
noptera). Iheringia Serie Zoologica 85: 133-136. 

Eisner, T. 1957. A comparative morphological study 
of proventriculus of ants (Hymenoptera, Formi- 
cidae). Bulletin of the Museum of Comparative Zo- 
ology at Harvard College 116: 437-490. 

Lappano, E. R. 1958. A morphological study of larval 
development in polymorphic all-worker broods 
of the army ant Eciton burchelli. Inscctes Sociaux 
5: 31-66. 

Nelson, J. A. 1924. Morphology of the honeybee lar- 
va. Journal of Agricultural Research 28: 1167-1229. 

Petralia, R. S. and S. B. Vinson. 1980. Internal anato- 
my of the fourth instar larva of the imported fire 
ant, Solenopsis iiivicta BUREN (Hymenoptera: 
Formicidae). Uiteruational journal of Insect Mor- 
phologi/ & Embryologif 9; 89-106. 

Tomotake, M. E. M. 1990. Morfologia comparada do tra- 
to digestivo de formigas em quatro trihos da subfam- 
ilia Ponerinae (Hi/menoptcra, Foniiicidae). Master 
Tesis— Rio Claro UNESP. 112p. 

Tomotake, M. E. M. 1996. Ultra-estrutura do proventri- 
culo de operdrins da subfamilia Poneriime (Hi//»t';;o/'- 
tera, Formicidae). PhD Tesis- Rio Claro UNESP. 
129p. 

Tomotake, M. E. M., F. H. Caetano and M. 1. Camar- 
go-Mathias. 1995a. The proventriculus ring: a 
comparison between Ponerinae and Dolichoder- 
inae ants subfamily (Hymenoptera, Formicidae). 
Acta Microscopica. 4: 318. 

Tomotake, M. E. M., F. J. Zara and F. H. Caetano. 
1995b. A musculatura externa do proventriculo 
de Neoponera villosa: Mudani^a da fase larval para 
a vida adulta. Proceedings of XII Eiwontro de Mir- 
mecologia, Sao Leopoldo. p. 99. 

Valentini, S. 1951. Sur L'adaptacion des larves de For- 
micoidea. Aniudes Des Sciences Naturalles et Zool- 
ogie 11: 249-276. 

Walker, J. R. and D. F. Clower. 1961. Morphology and 
histology of the alimentary canal of the imported 
fire ant queen {Sokmopsis saevissima ricliteri). An- 
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28. 

Wheeler, C. G and J. Wheeler. l'-»76. Ant larxae: re- 
view and synthesis. Proceedings of the Entomolog- 
ical Societi/ of Washington. 108pp. 

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Zara, F. J. and F. H. Caetano. 1998. Formaqao da 
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dae). Revista Brasileira de Biologia 58: 33-37. 



J. HYM. RES. 

Vol. 10(2), 2001, pp. 261-270 

Ultrastructure of Spermatozoa in Pleheia (Plebeia) droryana Friese 
(Hymenoptera: Apidae: Meliponina) 

Uyra Zama, Jose Lino-Neto, and Heiui Dolder 

(UZ, JL-N, HD) Dept. of Ceil Biology, State University of Campinas (UNICAMP), Campinas, 
Sao Paulo, Brazil; (JL-N) Dept. of General Biology, Federal University of Vic^osa (UFV), Vi^osa, 

Minas Gerais, Brazil 



Abstract. — In general, the spermatozoa of Plcbcia (Plcbcin) dwri/niui Friese are very similar to 
those described for other Hymenoptera. However, their arrangement in spermatodesmata bundles 
in the seminal vesicle has not yet been found in Apidae, this being a characteristic observed, to 
date, only in Symphyta, the Hymenoptera considered most primitive. The spermatozoa are long 
and thin, made up of a head connected to the tail at the position of the centriolar adjunct. The 
head includes an acrosomal vesicle, a perforatorium and a electron dense nucleus. The flagellum 
consists in a typical axoneme, two mitochondrial derivatives and two accessory bodies. Unlike 
most other Hymenoptera, the centriolar adjunct is very long and located between the nuclear base 
and the anterior extremity of the smaller mitochondrial derivative. It has recently been demon- 
strated that the structure and ultrastructure of hymenopteran spermatozoa are sufficiently varied 
so as to furnish consistent character matrices that can contribute to phylogenetic studies ("Sper- 
miocladistics")- Since no consensual phylogenetic hypothesis has yet been proposed for Apidae, 
the data presented here may be a contribution in this direction. 



The Apidae have been extensively stud- 
ied due to their economic and ecological 
importance, since they are pc^llinators, of- 
ten exclusively, of the majority of flower- 
ing plants, including species cultivated by 
man. The relation between these pollinat- 
ing agents and the plants they pollinate is 
so intimate that changes in the biodiver- 
sity of either group is certain to affect the 
other. The Apidae are also recognized as 
a diverse group with complex social be- 
haviour, which culniinate in advanced eu- 
social societies, a level observed only 
among Hymenoptera (a few bees anci 
wasps) and in the Isoptera. 

Within the Apidae, the tribe Apini, con- 
sisting in the subtribes Apina, Bombina, 
Euglossina and Meliponina {sciisii Roig- 
Alsina and Michener 1993), is particularly 
interesting because its members collective- 
ly display all levels of social behaviour. 
Ranging from solitary bees, as in some 
Euglossina, to advanced eusocial groups. 



such as the Apina and the Meliponina, 
passing though intermediate social behav- 
ior groupings as found in the Bombina 
and Euglossina. 

In spite of the unc^uestioned importance 
of the Apidae, so far neither morphologi- 
cal nor molecular studies have been able 
to establish an uncontested phylogeny for 
this group (Camargo and Pedro 1992b; 
Cameron 1991, 1993; Cameron et al. 1992). 
The establishment of the phylogeny of this 
group would undoubtedly be important 
for studies of the evolutionary mecha- 
nism, or mechanisms, leading to eust:>cial 
behaviour (Crozier and Pamilo 1996). 

Structural and ultrastructural character- 
istics of the spermatozoa, besides their 
own biological and taxonomic aspects, 
may be very interesting if this information 
can be used to form a character matrix for 
phylogenetic analysis. This information, 
associated with other character systems, 
could lead to a better understanding of the 



262 



Journal of Hymenoptera Research 



ev^olutionary relationships within the 
group ("spermiocladistics", Jamieson 
1987) as is being carried out for other an- 
imals, including insects (Baccetti 1972; 
Dallai 1979; Dallai and Afzelius 1990, 
1995; Carcupino et al. 1995; Jamieson et al. 
1999; Lino-Neto et al. 1999, 2000a, 2000b). 
The spermatozoal ultrastructure of only 
one apid species. Apis mellifera Limieaus, 
representing the Apini, has so far been 
studied in detail (Rothschild 1955; Hoage 
and Kessel 1968; Cruz-Holfing et al. 1970; 
Lensky et al. 1979; Woyke 1970; Lino-Neto 
et al. 2000b). Besides this species, in Me- 
liponina only some aspects of spermiogen- 
esis were investigated, including that of 
ScnpUotrigona pwstica Latreille (Cruz-Lan- 
dim and Beig 1980; Cruz-Landim et al. 
1980), MeUpowi quadrifasciata niifliidioiiics 
Lepeletier (Cruz-Landim et al. 1980; Cruz- 
Landim and Moraes 1980), Plcbciii {Plcbcia) 
droryaun Friese, FricscomcUtia {FriescouicUi- 
ta) varia Lepeletier, Lciirotrigoiia mucUcri 
Friese (Cruz-Landim et al. 1980). However 
these publications contain almost no in- 
formation on the mature sperm cell. 
Therefore, in this study, we characterize 
the structure and ultrastructure of Plcbcia 
{Plcbcia) droryana sperm sc~) as to furnish 
data that could be used iov future phylo- 
genetic research. 

MATERIAL AND METHODS 

Adult males of Plcbcia (Plcbcia) droryana 
were obtained from colonies maintained 
in the Central Apiary of the Federal Uni- 
versity of Vi^osa, MG, Brazil. 

Light Microscopy. — Seminal vesicles 
were dissected and broken open on clean 
glass microscope slicies, where the sperm 
were spread and fixed in a solution of 4'/i) 
(wt/vol) paraformaldehyde in 0.1 M phos- 
phate buffer, pH 7.2. After drying at room 
temperature, the preparations were ob- 
served with a photomicroscope (Olympus, 
BX60), equipped with phase contrast. 

To measure the nucleus, some of these 
preparations were stained for 15 min. with 
0.2 (xg/ml 4,6-diamino-2-phenylindole 



(DAPl) in phosphate buffered saline, 
washed, and mounted with Vectashield. 
They were examined with an epifluoresc- 
ence microscope (Olympus, BX60), 
equipped with a BP360-370 nm excitation 
filter. 

TrniisiJiissioii Electron Microscopy. — Sem- 
inal vesicles were dissected and fixed for 
3 hours in a solution containing 2.5% glu- 
taraldehyde, 0.2% picric acid, 3% sucrose 
and 5 mM CaCL in 0.1 M cacodylate buff- 
er, pH 7.2. The materials were post fixed 
in 1% osniium tetroxide, in the same buff- 
er, for 1-2 hours. Dehydration was carried 
out in acetone and embedded in Epon. Ul- 
trathin sections were stained with uranyl 
acetate and lead citrate and observed with 
the Zeiss LEO 906 transmission electron 
microscc^pe. 

RESULTS 

In the seminal vesicle, the spermatozoa 
of Plcbcia {Plcbcia) droryana are organized 
in spermatodesmata bundles, where the 
anterior region of the heads are embedded 
in a substance of medium electron density 
(Figs. 1, 2). The more central spermatozoa 
are situated slightly ahead of the lateral 
ones, so that a transverse section of this 
region shows acrosomes sectioned at dif- 
ferent levels (Fig. 2). However, some iso- 
lated spermatozoa also appear chaotically 
dispersed in the seminal vesicles (Figs. 5- 
7). 

The sperniatozoan of P. droryana is long 
and thin, measuring approximately 135 
|jLm in length (Fig. 3). The acrosome mea- 
suring about 1.2 (jtm and is made up of 
the acrosomal vesicle and the perforatori- 
um (Figs. 1, 5-6). The acrosomal vesicle is 
cone-shaped and covers the perforatorium 
along its entire length (Fig. 6). In trans- 
verse section, the acrosome is circular at 
the tip but beconies triangular, particular- 
ly the perforatorium towards the nucleus 
(Figs. 2, 8-9). Along the circular portion, 
an electron transparent layer ct^vers the 
perforatorium, separating it completely 
from the acrosomal vesicle. However, 



VoLUMi: 10, Number 2, 2001 



263 



when they are triangular this clear layer is 
reduced to patches at the vertices (Figs. 8- 
9). The perforatorium base penetrates 
about 70 nm into a small asymmetric cav- 
ity in the nuclear tip (Fig. 7). 

The nucleus measures approximately 
7.5 |jLm in length and is filled homoge- 
neously with dense chromatin. In trans- 
verse section, it is slightly oval, measuring 
approximately 0.18 |jLm in diameter at the 
anterior extremity and 0.45 jxm at the pos- 
terior (Figs. 2-7, 10-13). At the anterior tip 
there is a cavity in which the perforatori- 
um fits (Fig. 7), while posteriorly the nu- 
cleus tapers conically and is covered by 
thin electron transparent and electron 
dense material (Figs. 12-13, 15). 

The axoneme, measuring 126 [xm of 
length, presents the 9 + 9 + 2 pattern of mi- 
crotubules, with 9 single, external, acces- 
sory microtubules, nine doublets and a 
pair of single ones Lri the center of the ar- 
rangement (Figs. 18-21). In the first 0.28 
fxm, corresponding to the centriole, the ax- 
oneme consists only of the accessory mi- 
crotubules, the doublets and a dense 
amorphous substance (Fig. 16). The cen- 
tral microtubules begin posterior to the 
centriolar portion (Fig. 17). In the final 
portion, the axoneme is gradually disor- 
ganized, with the central microtubules 
and the nine doublets terminating first, si- 
multaneously, followed by the accessory 
microtubules (Figs. 21-24). 

The centriolar adjunct is very long, 
about 4.6 |jim in length, compact and elec- 
tron dense. It begins at the nuclear base 
and extends parallel to the axoneme until 
it fits onto the smaller mitochondrial de- 
rivative (Figs. 11-14). In longitudinal sec- 
tion, it has a rod-like shape while in trans- 
verse section it is approximately circular, 
with a diameter of about 0.2 ixm (Figs. 1, 
11-12, 14, 16-18). 

The mitochondrial derivatives are 
asymmetric in both length and diameter 
(Figs. 11, 13-14, 19). Anteriorly, the larger 
mitochondrial derivative begins next to 
the tapering nucleus (Fig. 13) and the 



smaller in contact with the posterior end 
of the centriolar adjunct. In transverse sec- 
tion, the derivatives are elipsoidal, with 
the larger one curving slightly over the 
smaller one (Fig. 19). Both have at least 
three regions: a dense material that fills in 
n"iost of the mitochondrial derivatives; a 
clear approximately central area and the 
region of the cristae, limited to that part 
of the periphery opposite the axoneme (a, 
b and c in Figs. 16-19). The large mito- 
chondrial derivative also has a region of 
regularly arranged paracrystalline mate- 
rial in the third that is most distal to the 
axoneme (p in Figs. 16-19). Anteriorly, the 
derivative extremities do not show any 
cristae (Figs. 13-14). 

The accessory bodies are located later- 
ally, between the axoneme and the mito- 
chondrial derivatives. In transverse sec- 
tions, they have a triangular shape (Figs. 
18-20). In the cenriolar adjunct region, 
there is only one accessory bociy present 
between the larger mitochondrial deriva- 
tive and the axoneme (Fig. 18). 

DISCUSSION 

The arrangement of spermatozoa in 
spermatociesmata observed in Plebcin ciwr- 
yana, has not been described for Apocrita. 
According to Quicke et al. (1992), this 
spermatozoa arrangement in bundles is 
characteristic of Symphyta, considered 
primitive Hymenoptera, in spite of some 
sheath fragments encountered by these 
authors in some Aculeata. The central 
spermatozoa of the sheaths are somewhat 
ahead of the others, as observed in P. dror- 
ynna, as also occurs in Xyeloidea and 
Phamphiloidea, which are considered the 
most basal Symphyta (Newman and 
Quicke 1999a). However, in the Siricidae, 
considered the family most closest to Acu- 
leata studied so far, the central spermato- 
zoa are inserted well ahead of the periph- 
eral ones, so that in transverse sections, 
they are observed in very different levels 
(Newman and Quicke 1999a). Although 
most of the spermatozoa are organized in 



264 



Journal of Hymenoptera Research 




Volume 10, Number 2, 2001 



265 



spermatodesmata in P. droiynnn, as in the 
symphytans, Trenicx sp. (Newman and 
Quicke 1999a) and Cnlameutn sp. (Quicke 
et al. 1992), some spermatozoa are free. 
Newman and Quicke (1999a) suggested 
that the observation of free spermatozoa 
in the seminal vesicle could be due to fix- 
ation or if they indicate a pre-transfer phe- 
nomenon. We believe that it is also pos- 
sible that these spermatozoa have either 
not yet been grouped into sperniatodes- 
mata, or even that not all spermatozoa are 
destined to became included in bundles. 

In all the apocritan non-Aculeata (par- 
asitic wasps) considered to date, the sper- 
matozoa appear isolated in the seminal 
vesicle, and no spermatodesmata frag- 
ments have been observed (ex. Quicke et 
al. 1992; Newman and Quicke 1998, 1999b; 
Lino-Neto et al. 1999, 2000a; Lino-Neto 
and Dolder 2001a, b). The fact that sper- 
matozoa organized in spermatodesmata 
occur in Symphyta and in at least one apo- 
critan Aculeata, which are considered, re- 
spectively, the most basal and the most 
derived hymenopteran groups, while not 
occurring in the apocritan non-Aculeata, is 
very intriguing. This suggests either that 
it could be a reversed character state in 
Aculeata or that this group derived di- 
rectly from the Symphyta, as is believed 
to have occurred with parasitic wasps. 
This latter hypothesis seenis less likely 



since morphological and molecular anal- 
yses suggest that Aculeata are the sister 
group of the Ichneumonoidea (Whitfield 
and Cameron 1998; Ronquist et al. 1999). 
The basic structure of the spermatozoa 
in P. drori/ana is quite similar to that de- 
scribed for other Hymenoptera, as well as 
for insects in general (Phillips 1970; Bac- 
cetti 1972). The acrosome of P. droryann, 
made up of an acrosomal vesicle and the 
perforatorium appears to be typical for 
Hymenoptera (Jamieson 1999), having 
been found in Symphyta (Quicke et al. 
1992; Newman and Quicke 1999a), in the 
Scelionidae, Trissolciis bnsnlis (Lino-Neto 
and Dolder 2000a), in Formicidae (Wheel- 
er et al. 1990) and in Apis nicllifera (Cruz- 
Hofling et al. 1970; Hoage and Kessel 
1968; Lensky et al. 1979; Peng et al. 1992, 
1993). In this last species, unlike the other 
Hymenoptera studied, the acrosome is al- 
most as long as the nucleus, measuring 
about 5.6 |jLm. The fact that the acrosome 
of P. droryann shows a circular cross sec- 
tion at the tip gradually being modified 
into a triangular form as it reaches to- 
wards the nucleus differs from other Hy- 
menoptera where this acrosome are al- 
ways circular (ex. Symphyta, Newman 
and Quicke 1999a; Cynipoitiea, Newman 
and Quicke 1999b; Chalcidoidea, Lino- 
Neto et al. 1999, 2000a; Lino-Neto and 
Dolder 2001b; Formicidae, Wheeler 1990), 



Figs. 1-14. Ultramicrographs of Plcbeia spermatozoa in seminal vesicle. 1-2, Longitudinal and transverse 
sections, respectively, of anterior region of a spermatodesm. 1, Acrosomal region (arrow) and portion of 
nucleus (n) embedded in less eletron dense extracellular material (*). The arrowhead indicates the centriolar 
adjunct. 2, Numbers 1-6 indicate acrosomes sectioned in anterior-posterior levels from tip to just above nu- 
cleus. 3-4, Phase contrast micrograph of a spermatozoa (3) and head region, DAPI-stained fluorescence of 
nucleus. The arrow indicates the head (h) and tail (t) limit. 5, Longitudinal section showing acrosome (ac) 
and nucleus. 6, Longitudinal section of acrosomal vesicle (av) and perforatorium (p). 7, Transition region of 
acrosome-nucleus showing perforatorium base fitting into cavity of nuclear tip. 8-10, Transverse section of 
acrosome tip (8), base of acrosome (9) and nucleus free of extra cellular material (10). 11-13, Longitudinal 
sections of nucleus-flagellum transition region. Arrows indicate connective material at nuclear base (12, 13); 
14, Longitudinal section at junction of centriolar adjunct and smaller mitochondrial derivative. Arrowhead 
indicates mitochondrial cristae. Abbreviations; n = nucleus; ac = acrosome; a\^ = acrossomal vesicle; p = 
perforatorium; ca = centriolar adjunct; ax = axoneme; M = larger mitochondrial derivative; m = smaller 
mitochondrial derivative. Scale bar: 1, 4, 8-9 = 3|jLm; 2 = 2|jL.m; 3 = 8|xm; 6 = 0,ljjLm; 7 = 0,2(a.m; 10-11, 14 
= 0,3|jLm and 5, 12-13 = 0,5|jLm. 



266 



Journal of Hymenoptera Research 




► ^ 



^ > 



20 / 







_ 22/ _ 23 




Figs. 15-24. Sequential transverse sections oi tlagella. 15, Nucleus-flagellum transition region. Arrow indi- 
cates material connecting nucleus to larger mitochondrial derivative. 16-17, Centriolar region of axoneme. 
Open arrow indicates first of central microtubules. 18-19, Sections of flagellum, at centriolar adjunct region 
and at both mitochondrial derivatives, respectively. The arrows indicate accessory bodies and (*) indicates 
central material between flagellar structures. 20-24, Final flagellar region. The nine doublets (arrowheads) and 
two central microtubules (small arrow) terminate first, followed by accessory ones (white arrowheads). Large 
arrows indicate accessory bodies. Abbreviations: a = less electron dense amorphous region; b = more electron 
dense amorphous region; c = cristae region; pc = paracristalline region in the larger mitochondrial derivative; 
ca = centriolar adjunct; ce = centriole; n = nucleus; ax = axoneme. Scale bar: 15-20 = 0,1 |xm; 21-22 = 0,06|jLm 
and 23-24 = 0,05(jLm. 



or maintains an oval cross section as in 
Apis mellifcra (q.v.) and Vespidae (personal 
observation). The acrosome of A. nielli fern 
also differs from that of P. drorymia due to 
the presence of a long anterior projection 
of the acrosomal vesicle (Cruz-Hofling et 
al. 1970; Hoage and Kessel 1968). The pen- 
etration of the perforatorium in the nucle- 
ar tip as occurs in P. droryana has been 
described for the majority of the hyme- 
nc-)pterans (ex. Quicke et al. 1992; Newman 



and Quicke 1999a; Wheeler et al. 1990). 
However, in Eurytomidae, Bcphrntclloidcs 
pomoruni Fabricius (Lino-Neto et al. 1999), 
and in the Pteromalidae, Nasonia vitripen- 
nis Walker (Hogge and King 1975), the 
perforatorium base is concave and has the 
same diameter as the nucleus in this re- 
gion, fitting directly ontti the anterior nu- 
clear surface, hi the majority of parasitic 
wasps, there is a third extracellular layer 
(the extracellular sheath), covering all of 



Volume 10, Number 2, 2001 



267 



the acrosome and extending along a vari- 
able length of the nucleus (Quicke et al. 
1992; Newman and Quicke 1999b; Quicke 
et al. 1992; Newman and Quicke 1998; 
Quicke et al. 1992; Lino-Neto et al. 1999, 
2000a; Lino-Neto and Dolder 2001b). Also, 
in some of these, the extra-cellular sheath 
gives rise to innumerable filaments, prob- 
ably representing a well developed gly- 
cocalix (Lino-Neto et al. 1999, 2000a; Lino- 
Neto and Dolder 2001b). 

In P. droryana, the nucleus is long, dense 
and usually appears homogeneously com- 
pacted. These characteristics are highly 
conserved in Hymenoptera, and the vari- 
ations observed have been in length and 
in the fact that this structure may be linear 
(ex. Quicke et al. 1992; Jamieson et al. 
1999; Wheeler et al. 1990; Lino-Neto et al. 
2000b), or twisted in a spiral, as in Chal- 
cidoidea (Lee and Wilkes 1965; Hogge and 
King 1975; Quicke et al. 1992; Lino-Neto 
et al. 1999, 2000a, 2000b; Lino-Neto and 
Dolder 2001b), Scelionidae (Lino-Neto and 
Dolder 2001a) and Diapriidae (Quicke, 
personal communicatit^n). The nucleus of 
P. droryiDin ends in a short cone, next to 
the anterior tip of the large mitochondrial 
derivative, and terminating in contact 
with the centriolar adjunct and axoneme. 
In Apis uieUifern, the final nuclear projec- 
tion is considerably longer and inserted in 
the axoneme, so that in cross section the 
nucleus is found surrounding the tips of 
the centriolar microtubules (Peng et al. 
1993; Lino-Neto et al. 2000b). In the ma- 
jority of the Hymenoptera, the nucleus is 
not tapered posteriorly but instead is 
abruptly truncated (Quicke et al. 1992; 
Newman and Quicke 1999a; Newman and 
Quicke 1999b; Quicke et al. 1992; Wheeler 
et al. 1990; Lino-Neto et al. 1999, 2000a; 
Lino-Neto and Dolder 2001b). 

The centriolar adjunct of P. drori/aiin is 
a well developed structure located par- 
allel to the axoneme and between the nu- 
cleus and the smaller mitochondrial de- 
rivative. This arrangement has also been 
found in some Symphyta (Newman and 



Quicke 1999a), Cynipoidea (Newman and 
Quicke 1999b), Ichneumonoidea (Quicke 
et al. 1998) and in A. mellifera (Lino-Neto 
et al. 2000b). However, in the Ichneumo- 
noidea this structure is comparatively 
short (Quicke et al. 1998) while in A. mel- 
lifera, it is extremely long, tapered ante- 
riorly, widening into a thick rod posteri- 
orly (Lino-Neto et al. 2000b). In the sym- 
phytan Treiiiex sp. (Newman and Quicke 
1999a) and in the Formicidae (Wheeler et 
al. 1990), the centriolar adjunct lies be- 
tween the nucleus and both mitochondri- 
al derivatives. On the other hand, in 
Chalcidoidea (Lino-Neto et al. 1999, 
2000a; Lino-Neto and Dolder 2001b) the 
centriolar adjunct is located laterally to 
the final portion of the nucleus, sur- 
rounding the nuclear-flagellum transition 
and extending parallel to the axoneme for 
a short distance, above the insertion of 
both mitochondrial derivatives. Contrary 
to the majority of these insects, no cen- 
triolar adjunct was encountered in Sce- 
lionidae (Lino-Neto and Dolder 2001a). 
The great variation in shape and location 
of the centriolar adjunct, differing from 
that known for most insects (Jamieson 
1982) is probably the reason for the ear- 
lier misinterpretations of this element in 
various Hymenoptera (Cruz-Hofling et 
al. 1970; Quicke et al. 1992). 

The mitochondrial derivatives of P. 
droryana are asymmetric not only in length 
but also in diameter. As a rule, the deriv- 
atives are straight (ex. Quicke et al. 1992; 
Jamieson et al. 1999; Wheeler et al. 1990; 
Lino-Neto et al. 2000b), but in Chalcidoi- 
dea (Lee and Wilkes 1965; Hogge and 
King 1975; Quicke et al. 1992; Quicke 1997; 
Lino-Neto et al. 1999, 2000a, 2000b; Lino- 
Neto and Dolder 2001b), Scelionidae 
(Lino-Neto and Dolder 2001a) and Diapri- 
idae (Quicke, personal communication) 
they spiral around the axoneme. The larg- 
er mitochondrial derivative beginning 
next to the final projection of the nucleus 
was also observed in A. ineUifera (Lino- 
Neto et al. 2000b) and in Cynipoidea 



268 



Journal of Hymenoptera Research 



(Newman and Quicke 1999b). This is not 
the case of the majority of the Hymenop- 
tera, where the larger mitochondrial de- 
rivative abuts the nuclear base, not over- 
lapping it (ex. Quicke et al. 1992; Wheeler 
et al. 1990; Jamieson et al. 1999; Newman 
and Quicke 1999a). In Megalyroidea 
(Newman and Quicke 2000), Diapriidae 
(Quicke, personal communication) and 
Scelionidae (Lino-Neto and Dolder 2001a) 
the large mitochondial derivative projects 
parallel to the nucleus for a considerable 
distance, and in this latter family, only one 
large mitochondrion is observed (Lino- 
Neto and Dolder 2001a). In transverse sec- 
tions of the P. don/aiin flagellum, four dis- 
tinct regions make up the larger derivative 
while only three are found in the smaller 
one. The same organization was observed 
in A. mdlifcra (Lino-Neto et al. 2000b) al- 
though Cruz-Hofling et al. (1970), Lensky 
et al. (1979) and Peng et al. (1992, 1993) 
have described the presence of paracrys- 
talline material also in the smaller deriv- 
ative. In the Formicidae the mitochondrial 
derivatives consist in three regions 
(Wheeler et al. 1990). However, the re- 
gions described in Formicidae are not 
analogous to those in the smaller deriva- 
tive of P. dronjiDui. In Formicidae, there is 
a clear area, a well developed region of 
cristae and the paracrystalline material sit- 
uated in the mitochondrion's first third, 
proximal to the axoneme (Wheeler et al. 
1990). Asymmetrical diameters of mito- 
chondrial derivatives are frequently 
found, occurring in the Symphyta (Quicke 
et al. 1992; Newman and Quicke 1999a), 
Cynipoidea (Quicke et al. 1992; Newman 
and Quicke 1999b), Megalyroidea (New- 
man and Quicke 2000) and Proctotrupo- 
idea (Quicke et al. 1992). However, bees 
are even more strongly asymmetrical 
(Cruz-Hofling et al. 1970; Hoage and Kes- 
sel 1968; Lensky et al. 1979; Peng et al. 
1992, 1993; Lino-Neto et al. 2000b). On the 
other hand, some Hymenoptera have 
symmetrical mitochondrial derivatives as 
in Formicidae (Wheeler et al. 1990) and 



Chalcidoidea (Lino-Neto et al. 1999, 
2000a). 

Plcbcia droryaun, as is common to most 
insects (Jamieson et al. 1999), has an axo- 
neme with the microtubules arranged par- 
allel to each other. This is not the case of 
Chalcidoidea (Lee and Wilkes 1965; Hog- 
ge and King 1975; Quicke et al. 1992; 
Quicke 1997; Lino-Neto et al. 1999, 2000a, 
2000b; Lino-Neto and Dolder 2001b), Sce- 
lionidae (Lino-Neto and Dolder 2001a) 
and Diapriidae (Quicke, personal com- 
munication) where they follow a spiraling 
course. Also in P. droryaun, the accessory 
microtubules are the last ones to terminate 
at the end of the axoneme. This character- 
istic is also observed in A. Dicllifcm (Peng 
et al. 1993; Lino-Neto et al. 2000b) and in 
Formicidae (Wheeler et al. 1990), while in 
Chalcidoidea (Lino-Neto et al. 1999; Lino- 
Neto and Dolder 2000a, b) and Ichneu- 
monoidea (Braconidae) (Newman and 
Quicke 1998) the accessory tubules termi- 
nate first. Unfortunately, this characteristic 
has not been taken in consideration by 
most studies of hymenopteran spemato- 
zoa. We believe this could be a useful pa- 
rameter to help separate the Aculeata, or 
parasitic wasps, from other Hymenoptera. 

The triangularly shaped accessory bod- 
ies, as found in transverse sections of P. 
droryaun, are encountered in most Hyme- 
noptera (Quicke et al. 1992; Jamieson et al. 
1999; Wheeler et al. 1990; Lino-Neto et al. 
2000b). They may be considerably reduced 
in Chalcidoidea (Lino-Neto et al. 1999, 
2000a; Lino-Neto and Dolder 2001b) and 
in the Scelionidae (Lino-Neto and Dolder 
2001a) so that, in some cases, they are dif- 
ficult to identify. The function of this 
structure has not been clearly established 
but they appear to be involved in the at- 
tachment of the mitochondrial derivatives 
on to the axoneme, since they do not occur 
between the centriolar adjunct and the ax- 
oneme. 

In P. droryaun a small central structure 
was identified between both the mito- 
chondrial derivatives and the axoneme 



Volume 10, Number 2, 2001 



269 



(see asterisk in Figs. 18 and 19). This struc- 
ture was initially described in Formicidae 
(Wheeler et al. 1990), but it is possible that 
it is present in the majority of Hymenop- 
tera (Lino-Neto et al. 2000b). 

Based on the characteristics compared 
above, the sperniatozoa of this bee are, for 
the most part, similar to the majority of 
the Hymenoptera (Jamieson et al. 1999). 
Some distinct differences stand out. For 
example: (1) the arrangement of sperma- 
tozoa in spermatodesmata in the seminal 
vesicle, (2) the presence of a very long cen- 
triolar adjunct between the nucleus and 
the smaller mitochondrial derivative and 
(3) the presence of abundant paracrystal- 
line material, exclusively in the large mi- 
tochondrial derivative. 

The identification of these characteris- 
tics and other more subtle ones suggest 
that the sperm cell can furnish a character 
matrix for Hymenoptera that will be use- 
ful for future phylogenetic studies. 

ACKNOWLEDGMENTS 

The authors would like to thank Professor Dr. Lu- 
cio Antonio de Oliveira Campos (DBG/UFV) for sup- 
plying the insects. This research was supported by 
the Brazilian Agencies CNPq and FAPESP. 

LITERATURE CITED 

Baccetti, B. 1972. Insect Sperm Cell. Advanced bisect 
Phifsiology 9: 315-397. 

Camargo, J. M. F. and S. R. M. Pedro. 1992a. System- 
atics, phylogeny and biogeography of the Meli- 
poninae (Hymenoptera, Apidae) a mini review. 
Apidohgic 23: 1-32. 

Camargo, J. M. F. and S. R. M. Pedro. 1992b. Siste- 
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J. HYM. RES. 
Vol. 10(2), 2001, pp. 271-279 

Seed-feeding Species of Megastigmtis (Hymenoptera; Torymidae) 
Associated with Anacardiaceae 

E. E. Grissell and G. L. Prinsloo 

(EEC) Systematic Entomology Laboratory, PSI, Agricultural Research Service, U. S. Department 

of Agriculture, c/o National Museum of Natural History, Washington, DC 20560-0168, USA; 

(GLP) Biosystematics Division, ARC-Plant Protection Research histitute, Private Bag X134, 

Pretoria 0001, South Africa 



Abstract. — Four species of Megastignnis Dalman have been reported as phytophagous in seeds 
of the plant family Anacardiaceae: M. pistaciae Walker, M. rhnsi (Hussey), M. thomseni (Hussey), 
and M. traiisvnaioisis (Hussey). All are Old World in origin. In this paper we summarize known 
host anci distribution data for each species, place Mcgastigiiius rinisi as a junior subjective synonym 
of Megastiginus transvaalensis, diagnose and illustrate all species, and provide a key to aid in the 
identification of females and, to some extent, males. Kiiown host-plant genera are: Pistacia L., Rhus 
L., Schiiuis L., Ozoroa Delile, and Lannca A. Richard. Megastignnis pistaciae and M. transvaalensis 
have been accidentally introduced into the New World. The former has long been established in 
California and is reported for the first time from Mexico; the latter is widely established, through 
host-shifting, in the states of Hawaii, California, Florida, and in Brazil and Argentina. 



The genus Megnstig)mis Dalman is rep- 
resented by 126 described species known 
from all geographic areas (Grissell 1999, 
Grissell and Heydon 1999) except possibly 
South America, w^here the only reported 
species appears to have been introduced 
(Perioto 1999, Grissell and Hobbs 2000). 
Species are most abundant in the Holarctic 
and Australian Regions. About one-third 
of Megastignnis species are phytophagous 
in seeds of 11 plant families, about one- 
third are parasitoids (or incjuilines) of gall- 
forming insects in the fruits, leaves, and 
stems of plants, and about one-third have 
no known hosts (Grissell 1999). 

Our paper focuses on a group of Mcgas- 
tigiiius species associated with the plant 
family Anacardiaceae. Although the fam- 
ily occurs worldwide, only four species of 
Mcgastigiiius have been reported from its 
seeds, and these all appear to be indige- 
nous to Africa, the Middle East, and the 
Mediterranean region. Introductions of 
two of these species have been reported 
for the New World: M. pistaciae Walker, 



introduced into California (see species sec- 
tion below), and M. transvaalensis (Hus- 
sey), found in the states of Hawaii, Cali- 
fornia, Florida, and recently in Brazil and 
Argentina (see species section below). 

Of the Mcgastigiiius species attacking 
Anacardiaceae, three have been recog- 
nized based as much upon their host pref- 
erence as upon their morphological dis- 
tinctness: Mcgastigiiius pistaciae reared 
from species of Pistacia L., Megastignnis 
rliusi (Hussey) reared frc^m species of Rhus 
L., and Megastignnis transvaalensis reared 
from species of Scliiinis L., a South Amer- 
ican tree introduced into Africa. The 
fourth species, Megastignnis thomseni (Hus- 
sey), was reared from a questionable host 
(now confirmed as Ozoroa Delile, see dis- 
cussion under thomseni) and has been 
known only from the type series since its 
description. With only one published ex- 
ception, whenever a species of Megastig- 
nnis has been identified from Pistacia, 
Rhus, or Schinus, the name given is always 
the one historically associated with the 



272 



Journal of Hymenoptera Research 



host plant. The exception was published 
by Furth (1985:166) who reported Mcgns- 
tignnts pistiicinc as the ". . . main pest of R. 
[Rhus] tripartita [(Ucria) DC)] fruits" and 
considered it to be an "oliphagous [sic]" 
wasp based on the previous Old World 
host records from Pistacia spp. (Davatchi 
1958, Romanenko 1972). Furth's speci- 
mens now reside in the National Museum 
of Natural History (Washington, DC) and, 
based on morphological criteria devel- 
oped in our studies, they are nc^t M. pis- 
taciac but are M. traiisvaalciisis as explained 
below. 

In this paper we discuss all species of 
Mcgastignius associated with seeds of An- 
acardiaceae, summarize all distribution 
and host data, and present a key. Our re- 
view is based on comprehensive new data 
for hosts and distributions based upon ex- 
tensive surveys undertaken by both au- 
thors in South Africa. Additonal Old 
World material was collected by Simeon 
van Noort in Western Cape Province, 
South Africa, and Alain Roques in the 
Mediterranean Region). Extensive New 
World data have been provided by the 
sampling of Stephen Hight (USDA, Forest 
Service, Volcano, Hawaii), Greg Wheeler 
(USDA, Agricultural Research Service, Ft. 
Lauderdale, Florida), and their colleagues, 
as well as Richard Rice (University of Cal- 
ifornia, Parlier, California). The material 
examined, in excess of 5000 specimens, is 
housed in the National Collection of In- 
sects, PPRl, Pretoria; the South African 
Museum, Cape Town; and the National 
Museum of Natural History, Washington, 
D.C. 

Special mention should be made con- 
cerning males of Megastigiuus species as- 
sociated with Anacardiaceae. In M. pista- 
ciae and M. trausvaalciisis males are ex- 
tremely variable in appearance (few males 
of M. thomseui are known). Small individ- 
uals are yellow with no modifications of 
the stigma (Fig. \) relative to females. 
Large males, however, have the head, 
lower mesosoma, and metasoma mostly 





Figs. 1-3. Mcgasti^jiiiis trausvaalciisis, stigmal area, 
male, showing variation. 1, Smallest specimens (all 
yellow). 2, Intermediate specimens (yellow with some 
black). 3, Largest specimens (black with some yel- 
low). 

black with yellow markings, the stigma is 
enlarged, and it is surrounded by a stig- 
mal cloud (Fig 3). If the two extremes 
found within males of both M. pistaciac 
and M. transvaalcusis (and presumably 
thomseui) were distinct they might repre- 
sent dimorphic male forms, however they 
are bridged by intermediate forms (Fig. 2). 
The condition does not seem to be poly- 
morphic, as there is a gradual cline be- 
tween extremes. Although some wing 
characters help distinguish females re- 



VoLUMi; 10, Number 2, 2001 



273 



12 14 16 18 20 22 24 



26 28 30 32 34 



Reared from Rhus spp. # 
Reared from Schinus spp. Q 
Host plant unknown A 




10 12 14 16 18 20 22 24 26 28 30 32 34 36 



Fig. 4. Distribution of Megastigmus tran^vaalcnsis in South Africa. Symbols indicate plant host genus from 
which specimens were reared or collected. 



gardless of size, they do not work for 
males. Small yellow males have slightly 
more setae on the wings than females, but 
larger black males are considerably more 
setose than even the small yellow males. 
In general, males of all species are rela- 
tively uncommon compared to females. 

At first glance the large black males 
would seem to bear no relationship to 
their conspecific females. That they are 
conspecific is provided by three lines of 
evidence. Extremes of males may be found 
in rearings froni the same host, locality, 
and time for both M. trnusvanlcusis and M. 
pistncinc. In the field we have seen small 
yellow males of M. transvnalcnsis mating 
with females only to have larger black 
males chase them away and initiate mat- 
ing themselves. In addition, DNA analysis 



shows no differentiation within male phe- 
notypes of M. trniisvnalcusis and females 
(Scheffer and Grissell in preparation). 

Megastigmus transvaalensis (Hussey) 

(Figs. 1-3, 4-7) 

Eiiinegastigiuus transvaalensis Hussey 1956:161- 
162 (Figs. lg,h, 4). Holotype fennale, Pretoria, 
Transvaal, South Africa (The Natural History 
Museum, London, examined); 5 female, 8 
male paratypes, same data as holotype (The 
Natural History Museum, London; "Hussey 
private collection"), reared from seeds of 
Sdiinus niollc L. 

Enmegastignnis rhusi Hussey 1956:161 (Figs. le,f, 
3). Holotype female, Bloemfontein, Orange 
Free State, South Africa (The Natural History 
Museum, London, examined); 2 female, 4 
male paratypes, same data as holotype (The 
Natural History Museum, London; "Hussey 



274 



Journal of Hymenoptera Research 




transvaalensis 





Figs. 5-11. Mc;^astis^iiius species, female. 5, 8, 10, Forewing setation on dorsal surface, except costal cell shows 
ventral setae only, b, 7, 9, 11, Heads, anterior view. 



Volume 10, Number 2, 2001 



275 



private collection"), reared from seeds of 
R}nis laiicca L. New synonymy. 

Megastigiiiiis triDisvaaleiisis: Boucek 1978:129. 
New combination from Euuiegnstigunis. 

Megastigmus rhiisi: Boucek 1978:129. New com- 
bination from EiDiicgastignius. 

Diagnostic Characters. — In both sexes of 
M. traiisvaalciisis the face (Fig. 6, 7) is nar- 
rowed and at most as wide as high (to 
slightly higher than wide); the least inter- 
ocular distance is greater than the scape 
length (excluding radicle); and the eye 
height is subequal to or greater than the 
least interocular distance. In some individ- 
uals the face is remarkably narrowed (Fig. 
6) accompanied by a similar lateral com- 
pression of the body. In females (but not 
males) the costal cell ventrally has at most 
a longitudinal row of setae in its apical 
third to half (Fig. 5; these setae are some- 
times broken off and the costal cell may 
appear asetose); the upper surface of the 
costal cell is bare even at its apical margin; 
the basal cell has no setae either niedially 
or along its posterior margin (i.e., the cu- 
bital setal line) (Fig. 5). The admarginal 
area of the forewing has setae extending 
as far (or nearly) as the stigmal vein (Fig. 
5). Females range from about 2 to 3 mm 
in body length (without ovipositor). 

Distribution. — In its natural range (based 
on association with Rhus), M. traiisvaalcii- 
sis is common throughout South Africa 
(Fig. 4) and has also been found in single 
collections from Zimbabwe, Kenya, Israel, 
and Morocco. 

Its putative areas of introduction (based 
on association with Sciiiuus) include Cali- 
fornia (first reported by Harper and Lock- 
wood 1961; reported also on introduced 
Rlius laiicca by Grissell and Hobbs 2000), 
Florida (first report by Habeck ct al. 1989), 
Hawaii (first report by Beardsley 1971 as 
Megastigmus sp.), Brazil (first report by 
Perioto 1999, as Megastigmus sp.), Argen- 
tina (S. Hight, pers. comm.). Reunion (Ha- 
beck ct al. 1989), Canary Islands (Grissell 
1979), and Mauritius (speciniens in Na- 
tional Collection of Insects, PPRI, Preto- 



ria). We have seen specimens from all 
these locations. 

Hosts. — The hosts of Megastigmus traiis- 
vaahmsis are now known to include spe- 
cies of Rhus and Schiiius, though the latter 
is a result of host shifting (Grissell and 
Hobbs 2000). Prior to our study, M. trans- 
vaalensis (as M. rfnisi) was reported only 
from Rhus lancea (Hussey 1956). As a re- 
sult of our studies, A/I. trausvaalensis is now 
known to be reared from seeds of R. an- 
giistifolia L., R. deutata Thunb., R. discolor 
E. Mey. ex Sond., R. laevigata laevigata L., 
R. laevigata villosa (L.) R. Fernandes, R. lu- 
cida L., R. magalismoutaua Sond., and R. 
zcyheri Sond. (all South African in distri- 
bution); R. vulgaris Meikle and R. natalen- 
sis Bernli. ex Kraus (Kenya); and R. tripar- 
tita (Ucria) DC (Morocco, Israel). Addi- 
tionally, adult M. trausvaalensis were col- 
lected on the following Rhus species, 
which may be host plants as well: R. chir- 
indensis Bak. (seeds were collected for 
rearing, but no wasps emerged), R. crosa 
Thunb., R. pyroides Burch., and R. rehman- 
uiana glabrata R. & A. Fernandes and R. 
pendulina Jacq. growing together (seeds of 
the former were collected for rearing, but 
no wasps enierged; the latter was in 
bloom only). 

Megastigmus trausvaalensis has also been 
reared from seeds of Schiuiis molle and S. 
tcrebinthifolius Raddi, though these are 
non-native hosts. 

Discussion. — In the past any specimen 
reareci from Schinus seed was considered 
to be M. trausvaalensis and any from Rhus 
seed was considered to be M. rliusi. A re- 
cent paper by Grissell and Hobbs (2000) 
presented the reasons for considering both 
to be conspecific and additional molecular 
analysis (Scheffer and Grissell, in prepa- 
ration) has confirmed this to be the case. 
Neither paper presented formal nomen- 
clatural decisions, and in the current pa- 
per we formally place M. rhusi as a junior 
subjective synonym of M. trausvaalensis. 
We choose the name trausvaalensis because 



276 



Journal of Hymenoptera Research 



it is the only name that has appeared in 
the literature since its publication. 

A single population was found in Ken- 
ya (collected by R. Copeland, specimens in 
USNM and Texas A & M University) in 
which all individuals (n = 90) appear to 
be extremely laterally compressed and the 
head appears as in Figure 7 as compared 
to the typical head as in Figure 6. Within 
n^iany other populations examined, similar 
appearing individuals were seen, but not 
the entire population. It is possible that 
the compressed condition is related to 
host seed morphology, or that a second 
species is present. So tar, DNA studies of 
the two forms have proven inconclusive. 

Mcgiistis^mus tniiisvanlciisis is believed to 
have been introduced into California 
along with its native host plant, RJiiis hiii- 
cea, and thereupon switched to introduced 
ornamental trees of the genus Schinus 
(Grissell and Hobbs 2000). Its presence in 
Hawaii, Florida, Brazil, and Argentina is 
presently under study by EEG and col- 
leagues who are attempting to trace its 
pattern of geographic movement (Scheffer 
and Grissell, in preparation). 

Megastigmiis pistaciae Walker 

(Figs. 8, 9) 

"Megastigmiis pistacinc Haliday": Walker 1869: 
313. [Noiiicii inidiini] 

Megastigmiis pistaeine Walker 1871:35. Syntype 
females, "S. France" and "Tuscany" (The 
Natural History Museum, London, exam- 
ined), taken on "Pistacia lentiseiis [L.] and on 
P. terehinthiis [L.]." 

Trogocarpiis baHestrerii Rondani 1877:204-205 
(Figs. 109-114). Lectotype female (designated 
by Bouc'cek 1974:245), Italy (La Specola, 
Florence), reared from seeds of Pistacia -ocra 
L. [Synonymized by Masi 1934:210.] 

Diagnosis. — In both sexes of M. pistnciiic 
the face is transverse, being wider than 
high, and the least interocular width is 
less than the eye height (Fig. 9). In females 
(but not males) the costal cell has 3 or 4 
ventral rows of short setae in its apical 
half or more (Fig. 8); the cubital setal line 



has at least a few setae along the posterior 
margin of the basal cell (Fig. 8); and the 
admarginal area has setae reaching to the 
stigmal vein. This is the largest species of 
Mcgnstigi}ius associated with Anacardi- 
aceae, with females reaching 3.5 mm to 5 
mm in body length (without ovipositor). 

Distribution. — This species is endemic to 
the Old World, with a known distribution 
in coastal Mediterranean areas (Boucek 
1977) from Italy and Greece where it ex- 
tends into Iran (Roques and Skrzypczyn- 
ska in preparation), the Crimea, Transcau- 
casia, and Turkmenia (Boucek 1977). It has 
been introduced into the United States 
where it occurs in California (Robinson 
1968), and we have seen specimens from 
Saltillo, Coahuila, Mexico (ex Pistacia sp.), 
which represent the first records for that 
country. We have seen one specimen in- 
tercepted at an American port of entry 
from Australia {Pistacia cJiiiicnsis Bunge) in 
1970 (National Museum of Natural Fiis- 
tory, Washington, DC). If the data are au- 
thentic, this is the only report of the spe- 
cies from Australia. 

Hosts. — Megastigmiis pistaciae is confined 
to seeds of species of the genus Pistacia. 
The following hosts have been reported 
for the natural range of M. pistaciae (only 
first report of hosts are cited): Pistacia vera 
and P. terehinthiis (De Stefani 1917), P. inii- 
tica Fisch. & C. A. Mey [now = P. atlantica 
niiitica (Fisch. & C. A. Mey) Rech. F.] (Ni- 
kol'skaya 1935), and P. atlantica Desf. 
(Davatchi 1958). In its introduced range 
(California), M. pistaciae has been reported 
attacking the following species of Pistacia: 
P. cJiinensis Bunge, P. integerrinia }. Stewart 
[now = P. cliinensis integerrinia (J. Stewart) 
Rech. F.], P. atlantica, P. lentiseiis, P. vera 
'Kerman' (commercial cultivar), and a hy- 
brid between P. atlantica and P. vera (Rice 
and Michailides 1988). 

Furth (1985) reported this species in as- 
sociation with rearings from Rhus tripartita 
(in Israel), but based on an examination of 
voucher specimens in the National Muse- 



Volume 10, Number 2, 2001 



277 



um of Natural History, Washington, DC, 
these specimens are M. trniisvaiilcnsis. 

Biology. — The biology of this species (as 
bnllestrerii) was discussed by De Stefani 
(1908, 1917). In Italy, eggs were deposited 
in June and July. Larvae were mature by 
September and overwintered until May 
when they pupated and emerged. Gener- 
ally there was only a single generation per 
year, but occasionally a few of the larvae 
pupated in August and September. At this 
time most of the host seeds were too hard 
for oviposition, but a few soft young fruits 
could be found and two generations per 
year occurred. Similar findings were made 
in Greece (Agnostopoulos 1938, as ballcs- 
trerii), Tunisia (Jarraya and Bernard 1971), 
and California (Rice and Michailides 
1988). Zerova and Seryogina (1994) illus- 
trated the damaged seed. Males of this 
species are reportedly uncommon, rang- 
ing only up to about 4% (Rice and Mi- 
chailides 1988). Additional observations 
for this species in California are given by 
Vettel and Harper (1969), Wiard (1969), 
and Rice and Jones (1996). 

Pnrasitoids. — Two eurytomids, Si/copJiiln 
bigiittntd (Swederus) and Eurykvua rosac 
Nees, have been reported as parasites of 
M. pistacinc (Davatchi 1958) in its endemic 
range. 

Discussion. — Walker (1869) created a iio- 
nicii iiudiiiu when he published the name 
"Mcgnstiginus pistacinc Haliday." Haliday 
neither used the name nor described the 
species. Walker (1871) eventually de- 
scribed the species. Boucek (1974) dis- 
cussed the types of bnllestrerii and the 
question of synonymy, which might be at- 
tributed either to Masi (1934) by implica- 
tion or directly to Nikol'skaya (1935:83). 
Nikol'skaya (1935) redescribed and illus- 
trated the female (as pistncinc) as did Zer- 
ova and Seryogina (1994) and Roques and 
Skrzypczynska (in preparation). 

Megastigmtis tlwmseni (Hussey) 

(Figs. 10, 11) 

Euniegasfigmus ihomseni Hussey 1956:159-161 
(Figs. lc,d). Holotype female, Wolhuitensen- 



skop (misspelling for Wolhuterskop), Trans- 
vaal, South Africa (The Natural History Mu- 
seum, London, examined); 4 female, 2 male 
paratypes same data as holotype (The Nat- 
ural History Museum, London; "Hussey pri- 
vate collection"). 
Megastigmus tlwmseni: Boucek 1978:129. New^ 
combination from Euiuc;,^astigiuus. 

Dingiiosis. — In both sexes of M. tlioiuseni 
the face is transverse, being wider than 
high, and the least interocular width (Fig. 
11) is greater than the eye height. In fe- 
males (but not males) the costal cell ven- 
trally has at most a median row of setae 
in its apical third to half (Fig. 10; some- 
times these setae are broken off and the 
ctistal cell appears asetose); the cubital se- 
tal line has no setae along the posterior 
margin of the basal cell (Fig. 10); and the 
area proximal (imier) to the stigma and 
the admarginal area are essentially aseto- 
se. Females of this species range from 3 to 
4 mm in body length (excluding oviposi- 
tor). 

Distribution. — Apart from the type lo- 
cality, this species is also reported here 
from the following South African locali- 
ties: Rustenburg and Broederstroom 
(North-West Province), Pretoria (Gauteng 
Province), Thabazimbi (Northern Prov- 
ince), Lake St. Lucia (Kwazulu-Natal 
Province, and Richtersveld (Northern 
Cape Province). It also has been collected 
from a single locality in Kenya (Coast 
Province, El Nino road to Mica Creek, 
May 13, 1999, R. Copeland). 

Hosts. — The types were reared from 
seeds of "witharpuisbos", a common 
name that Hussey suggested might refer 
to Heerin sp. Based on our knowledge of 
inciigenous species, the common name ac- 
tually applies to Ozoron pnniculosn (Sond.) 
R. and A. Fernandes, from which thoiiiseiii 
has subsequently been reared. This should 
be considered the correct host. We also 
have seen specimens reared from Ozoron 
obovntn (Oliv.) R. & A. Fernandes, O. pnn- 
iculosn, and Lnnnen discolor (Sond.) Engl. 

D/sc?/ss/o/;.— Boucek (1978:129) trans- 



278 



Journal of Hymenoptera Research 



ferred this species to the genus Megnstig- 
miis. It has remained unknown since its 



description, and its host is herein positive- 
ly identified for the first time. 



KEY TO SPECIES OF MEGASTIGMUS ATTACKING SEEDS OF ANACARDIACEAE 

1 Females and males: Face transverse, wider than high; eye height slightly to much less than least 
interocular distance (Figs. 9, 11) 2 

- Females aiid males: Face about as high as wide; eye height equal to, or greater than, least 
interocular distance (Figs. 6, 7) transvaalensis (Hussey) 

2 Female: Forewing with costal cell ventrally with at most a single row of setae in apical 1/2 
midway between front edge and submarginal vein (Fig. 10); admarginal area asetose (Fig. 10); 
basal cell, basal vein, and cubital setal line at most with 3 or 4 setae; only last segment of club 
with ventral micropilose area. (Small yellow males same for basal cell and area; large black 
males not yet known for this species) thomseni (Hussey) 

- Female: Forewing with costal cell ventrally with 3 or 4 rows of setae in apical 1/2 to 2/3 (Fig. 
8); admarginal area with several setae extending as far (or nearly) as stigmal vein (Fig. 8); basal 
cell, basal vein, and cubital setal line with more than 5 setae; last 2 segments of club with 
ventral micropilose area. (Small yellow males as for female; large black male: costal cell ventrally 
densely setose, upper surface with setae in apical 1/3) pistaciae Walker 



Acknowledgments 

During the course of this study we were given as- 
sistance in our field work and in musuem research 
by the following individuals for whom we express 
our sincerest thanks: Simon van Noort, Vincent 
Whitehead, and Hamish Robertson, South African 
Museum, Cape Town; David McDonald and John 
Donaldson, National Botanic Institute, Kirstenbosch, 
Cape Town; Jan Giliomee and Henk Geertsema, Uni- 
versity of Stellenbosch, Stellenbosch; Brian Barnes, 
ARC-Fruit, Vine and Wine Research Institute, Stellen- 
bosch; Gail Littlejohn and Mark Wright, Agricultural 
Research Council, Elsenburg; John Marshall, De Muel 
Farm, Ceres; Percy Watkins, Green Valley Nuts, 
Prieska; John Heraty, University of California, Riv- 
erside, California; Chris Desjardins, University of 
Maryland, College Park, Maryland; Richard Rice, 
University of California (Davis Campus), Pariier, Cal- 
ifornia; Nelson Perioto, Instituto Biologico, Ribeirao 
Preto, Brazil; Gregory Wheeler, USDA, ARS, Ft. Lau- 
derdale, Florida; Stephen Hight, USDA, FS, Volcano, 
Hawaii; R. Copeland and R. Wharton, Texas A & M 
University, College Station, Texas, with the aid of a 
USAID-sponsored fruit sampling program, ICIPE, 
Kenya; Alain Roques, INRA-Zoologie Forestiere, 
Olivet, France; and Sonja Scheffer, Systematic Ento- 
mology Laboratory (SEL), USDA, ARS, Beitsville, 
Maryland. We are also grateful to John Noyes for the 
loan of type material from the Natural History Mu- 
seum, London, and to him, Thomas Henry (SEL), Da- 
\id Smith (SEL), and two anonymous reviewers for 
reading the manuscript and making many helpful 
suggestions. 



Literature Cited 

Agnostopoulos, P. T. 1938. [Pests of hazel, chestnut 
and pistachio]. Dciidrokroiologia Eiircna 3: 497- 
544. [In Greek.] 

Beardsley, J. W. 1971. Notes and Exhibits: Mcgastig- 
nuis sp. Proceedings^ of the Eiitoiiiological Society of 
Hawaii 2\: 28. 

Boucek, Z. 1974. On the Chalcidoidea described by C. 
Rondani. Redia 55: 241-285. 

Boucek, Z. 1977. A faunistic review of the Yugosla- 
vian Chalcidoidea (Parasitic Hymenoptera). Acta 
Entomologica Jiigoslavica 13 (SuppL): 1-145. 

Boucek, Z. 1978. A study of the non-podagrionine To- 
rymidae with enlarged hind femora, with a key 
to the African genera, journal of the Eiitoiiiologicnl 
Society of Southern Africa 41: 91-134. 

Davatchi, G. A. 1958. Etude biologique de faune en- 
tomologique des Pistacia sauvages et cultives. Re- 
vue de Patliologie Vi'gctalc et d'Entoniologie Agricole 
de France 37:3-166. 

De Stefani, T. 1908. L'In>ctto dei frutti de pistacchio e 
inodo di liniitarne i danni. Istituto di Zoologia e 
Anatomia Comparata della R. Universita, Paler- 
mo. 61 pp. 

De Stefani, T. 1917. [Megastigitnis ballestrerii, a hyme- 
nopteron living on pistacio tree and turpentine 
tree in Sicily.] Bolletiiio Studi Inform R. Giodano 
Coloniale di Palermo 4: 101-131. [In Italian. | 

Furth, D. B. 1985. The natural history of a sumac tree, 
with an emphasis on the entomofauna. Traiisac- 
tioiifi of tlie CoiDiecticut Academy of Arts and Sci- 
ences 46:137-234. 

Grissell, E. E. 1979. Torymidae, pp. 748-769. /;;, K. V. 



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Krombein, P. D. Hurd, D. R. Smith, and B. D. 
Burks editors, Catalog of Hi/iiit'iioptcra in Aiiicricn 
Nortfi of Mexico. Vol 1. Symphyia and Apocrita, in 
Washington, D.C.: Smithsonian Institution Press. 
1198 p^T 

Grissell, E. E. 1999. An annotated catalog of world 
Megastigminae (Hymenoptera: Chalcidoidea: 
Torymidae). Coiitributiom; of the American Ento- 
mological Institute 31(4): 1-92. 

Grissell, E. E. and S. Heydon. 1999. The identity of 
two unplaced New World Megastigminae (Hy- 
menoptera: Torymidae). Proceedings of the E)ito- 
niological Societi/ of Washington 101: 611-613. 

Grissell, E. E. and K. R. Hobbs. 2000. Megastigmus 
transvaalensis (Hussey) (Hymenoptera: Torymi- 
dae) in California: Methods of introduction and 
evidence of host shifting, pp. 265-278. /;;, A. D. 
Austin and M. Dowton (Eds) The Hi/nienoptera: 
Evolution, Biodiversity and Biological Control. CSI- 
RO Publishing, Melbourne, Australia. 

Habeck, D. H., F. D. Bemiett, and E. E. Grissell. 198'-*. 
First record of a phytophagous seed chalcid from 
Brazilian peppertree in Florida. Florida Entomol- 
ogist 72:378-379. 

Harper, R. W. and S. Lockwood. 1961. Bureau of En- 
tomology. Forty-first Annual Report. California 
Department of Agriculture Bulletin 2:127-129. 

Hussey, N. W. 1956. A new genus of African Megas- 
tigminae (Hymenoptera: Chalcidoidea). Proceed- 
ings of the Royal Entomological Society ofLondoji (B) 
25:157162. 

Jarraya, A., and J. Bernard. 1971. Premieres observa- 
tions bioecologiques sur Megastignnis pistaciae en 
Tunisie. Aiuiales de I'Institut National de la Reclier- 
che Agro}iomique le Tunisie 44: 1-28. 

Masi, L. 1934. Nota sur Calcididi dell'Isola di Rodi. 
Bollettino delta Societa Entomologica Italiana 66: 210. 

Nikol'skaya, M. N. 1935. [Pistacia seed-eating chalci- 
dids and their parasites (Hvmenoptera, Chalci- 
didae)]. Plant Protection, Lenijigrad 1935: 81-87. 
[In Russian.] 

Perioto, N. W. 1999. [First record of the genus Megas- 
tignnis Dalman, 1820 (Hymenoptera: Torymidae) 
and first record for the subfamily Megastigminae 



from Brazil.] Arquivos do Instituto Biologico Siio 
Paulo 64 (1997): 115-116. [In Portuguese.'] 

Rice, R. E. and R. Jones. 1996. Seasonal monitoring of 
the pistachio seed chalcid. Kearney Plant Protec- 
tion Group, Plant Protection Quarterly 6(l):l-3. 

Rice, R. E. and T. J. Michailides. 1988. Pistachio seed 
chalcid, Megastigmus pistaciae Walker (Hymenop- 
tera: Torymidae), in California. Journal of Econom- 
ic Entomology 81:1446-1449. 

Robinson, D. W. 1968. California Department of Agri- 
culture pistachio seed chalcid progress report 6S-1. 2 
pp. 

Romanenko, K. E. 1972. [The principal pests of fruit 
of pistachio (Pistacia) and the possibilities for 
their biological or chemical control in Kirgiza]. 
Proceedi)igs of the 13th International Co)igress of En- 
tomology 3:85 [In Russian.] 

Rondani, C. 1877. Vesparia parasita non vel minus 
cognita. Bollettino delta Societa Entomologica Itali- 
ana 9: 166-213. 

Roe]ues, A. and M. Skrzypczynska. [In prep.] Seed- 
infesting chalcids of the genus Megastigmus Dal- 
man (Hymenoptera: Torymidae) native and in- 
troduced to Europe: taxonomy, host specificity 
and distribution, journal of Natural History. 

Vettel, W. G. and R. W. Harper. 1969. California De- 
partment of Agriculture Pistachio Seed Chalcid Pro- 
gress Report No. 69-1. 

Walker, F. 1869. Notes on Chalcididae; and descrip- 
tions of a new species of Megastignnis. Transac- 
tions of the Royal Entomological Society of London 
1869: 313-314.' 

Walker, F. 1871. Notes on Chalcid iae. Part U.-Eurytom- 
idae and Torymidae, pp. 19-36. London: E. W. 
Newman. 

Wiard, W. W. 1969. Observations on the newly intro- 
duced pistachio seed chalcid, Megastigmus pista- 
ciae Walker. Califoriiia Department of Agriculture 
Report jaiuiary 1969. 

Zerova, M. D., and L. Y. Seryogina. 1994. [The seed- 
feeding Chalcidoidea of Palaearctics]. Kie\': National 
Academy of Sciences of Ukraine, L. L. Schmal- 
hausen Institute of Zoologv, Naukt^xa Dumka. 
237 pp. [In Russian.] 



J. HYM. RES. 

Vol. 10(2), 2001, pp. 280-281 



NOTE 



Polynema Haliday, 1833 (Insecta, Hymenoptera): Designation of 
Polynema flavipes Walker, 1846, as the Type Species 

John T. Huber and Z. Boucek 

(JTH) Canadian Forest Service, Natural Resources Canada, c/o ECORC, K. W. Neatby Building, 

C. E. F., Ottawa, ON, KIA 0C6, Canada; (ZB) c/o Department of Entomology, The Natural 

History Museum, Cromwell Road, London, SW7 5BD, England 



Abstract. — Polyucma flavipes Walker, 1846 (= P. oviiloniiii Haliday, 1833, misidentification oi Icli- 
nciiiiioii ovulonini Linnaeus) is designated as type species of Polynema, a widespread genus of 
Mymaridae. 



For more than 50 years it has been 
known that the previously clesignateci 
type species of the widespread genus Po- 
h/)iciiiii (Mymaridae, Fiymenoptera) need- 
ed to be fixed by the ICZN. This is done 
here, using the first of the five species 
originally included in Poh/ncnin by HaVi- 
day, namely P. flavipes, but initially mis- 
identified by him as P. ovulorum Linnaeus, 
1758. 

The genus Poh/iicuin was briefly de- 
scribed by Haliday (1833: 269). In the sec- 
ond part of his paper (1833: 347-348) he 
included five species in his genus: Ichucii- 
inou oviiloruni Linnaeus and four new 
ones. The former species was quoted later 
by Westwood (1839: 78) in his "examples 
of species of the British genera" as a "typ- 
ical species" of the genus Polyiicnin, an ac- 
tion that was later generally understood as 
a formal designation of the type species of 
genera, as ruled in ICZN Opinion 71 
(1922). 

Soon after his proposal of Poli/iiciiin, 
Haliday himself discovered that the Lin- 
nean species ovulonii}! could not belong to 
Polynomi, not even to Mymaridae, because 
he (Haliday) had misidentified the species. 
At that time, Haliday passed many of his 
chalcidological notes on to Francis Walker 



who eventually published them, as 
proved by his statement (1846: 49) "The 
following descriptions are, excepting a 
few additions, extracted from MSS kindly 
given to me by Mr. Haliday." It was there- 
fore Walker (1846: 52) who gave a new 
name, Poh/ncinn flavipes, to the species that 
Haliday had earlier misidentifieci, and 
added the comment "oviilonim olim; no- 
men errore ortum" [earlier ovuloriDii, the 
name used by mistake]. Despite this, how- 
ever, the error was repeated several times 
before 1960, lastly in the important work 
by Debauche (1948), who redescribed (pp. 
212-213) the species in c][uestion as Pohj- 
iieiua ovulorum (L.). Debauche (1949: 6, 7) 
and Soyka (1956: 2, 3) further discussed 
the problem but without a satisfactory res- 
olution, and Soyka (1956: 76) redescribed 
what he thought was Poh/)ieiun ovuloruui, 
based on a specimen collected by him in 
Austria in 1944 that he incorrectly desig- 
nated as lectotype and genotype. 

Another slight confusion was due to 
Hincks (1950: 177) who also referred to the 
misidentification problem. Hincks stated 
that the genotype [of Poh/)ienm] is the 
same as Eutrielie gnieilis Nees, 1834 ( = 
ovulorum Haliday nee Linnaeus). This was 
accepted by Mathot (1968: 276), who also 



Volume 10, Number 2, 2001 



281 



referred to the problem. Graham (1973) 
found and examined one original female 
specimen of Eiitridic gracilis Nees, labelled 
it as the lectotype, and showed that it is a 
species of Pohjncina not identical with P. 
flavipcs Walker (ovuloruni sensu Haliday). 

The type material of IcIniciDiioii ovuloruni 
could not be examined because it has been 
long lost, but the recent concensus is that 
it belonged to the present family Scelion- 
idae (superfamily Platygastroidea) (Bou- 
cek 1981: 18; Graham 1982: 228-229; John- 
son 1992: 605, and references therein). 

The identity of Poh/neuin flavipcs Walker, 
a replacement name for "ovuloruni sensu 
Haliday, 1833", was objectively defined by 
Hincks (1950: 181-183) who designated a 
lectotype (in the Haliday collection, Dub- 
lin) for the species. Graham (1982: 229) 
also examined the lectotype and con- 
firmed that it belongs to the present valid 
genus Polynenia Haliday. He suggested 
that the logical course would be to inval- 
idate Westwood's designation of Ichiieu- 
nioii ovuloruni Linnaeus as type species of 
Poli/nenin and to designate Polynenia flavi- 
pcs Walker, 1846 ( = P. ovuloruni Haliday, 
1833, misidentification of Ichneumon ovu- 
loruni Linnaeus) as type species of Poly- 
nenia. We concur with Graham's sugges- 
tion. Hence, under ICZN (1999) Article 
70.3.2 (4th edition, valid from 1st January, 
2000), we herewith fix Pohfiienia flavipcs 
Walker, 1846, misidentified as Polynenia 
ovuloruni (Linnaeus, 1758) by Haliday, 
1833, as type species of the genus Polyne- 
nia Haliday, 1833. 

LITERATURE CITED 

Boucek, Z. 1981. A biological solution to the identity 
of a Linnean chalcid wasp (Hymenoptera). £;;- 
tomologisi's Gazette 32: 18-20. 



Debauche, H.R. 1948. Etude sur les Mymarommidae 
et les Mymaridae de la Belgique (Hymenoptera 
Chalcidoidea). Meinoircs dii Miiscc Roi/al d'Histoirc 
Nntuirlle de Belgique 108: 1-248. 

Debauche, H.R. 1949. Exploration du Pare National 
Albert, Mission G.F. de Witte (1933-1935). 49: 1- 
105 + 13 plates. 

Graham, M.W.R. de V. 1973. The identity of Eiitriche 
graeilis Nees (Hymenoptera: Mymaridae). Ento- 
iiiologist's Gazette 24: 362-364 

Graham, M.W.R. de V. 1982. The Haliday collection 
of Mymaridae (Insecta, Hymenoptera, Chalcidoi- 
dea) with taxonomic notes on some material in 
other collections. Proceedings of the Roi/al Irish 
Academy, B 82: 189-243. 

Haliday, A.H. 1833. Essay on the classification of the 
parasitic Hymenoptera of Britain, which corre- 
spond with the Iclineumones minuti of Linnaeus. 
Entomological Magazine 1: 259-276, 333-350. 

Hincks, W.D. 1950. Notes on some British Mymaridae 
(Hym.). Transactions of the Society for British En- 
tomology 10: 167-207. 

ICZN. 1999. lnter)iational Code of Zoological Nomencla- 
ture. Fourth Edition. The International Trust for 
Zoological Nomenclature, c/o The Natural His- 
tory Museum, London. 306 pp. 

ICZN Opinion 71 (1922) Interpretation of the expres- 
sion "typical species" in Westwood's (1840) Syn- 
opsis. Smithsoiuan Miscellaneous Collectio)is 73(1): 
16-18. 

Johnson, N.F. 1992. Catalog of world species of Froc- 
totrupoidea, exclusive of Platygastridae (Hyme- 
noptera). Memoirs of the American Entomological 
Institute no. 51. 825 pp. 

Linnaeus, C. 1758. Si/stema naturae per reg)ia triad na- 
turae, secundum classes, ordines, genera, species, cum 
characteribus differcutiis, sy)ionymis, locis. Tomus I. 
Holmiae, Sweden. 824 pp. 

Soyka, W. 1956. Monographie der Polynemagruppe. 
Abhandlungen der Zoologisch-Botanischen Geselles- 
chaft in Wien 29: 1-115. 

Mathot, G. 1968. Mymaridae nouveau d'Afrique cen- 
tral (Hymenoptera: Chalcidoidea). Revue de Zool- 
ogie et de Botanique Africaifws 78: 265-276. 

Walker, F. 1846. Descriptions of the Mymaridae. An- 
nals and Magazine of Natural History 18: 49-54. 

Westwood, J.O. [June] 1839. Pp. 78-79 in: Synopsis of 
the geiwra of British insects: 49-80 [Issued with An 
introduction to the modern classification of in- 
sects. London.] 



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CONTENTS 
(Continued from front cover) 



SCHMIDT, S. and G. A. P. GIBSON. A new species of the genus Orussonia Riek and the 

female of O. depressa Riek (Hymenoptera: Symphyta, Orussidae) 113 

SHAW, S. R., G. SALERNO, S. COLAZZA, and E. PERI. First record of Aridehis rufotes- 
taceus Tobias (Hymenoptera: Braconidae, Euphorinae) parasitizing Nezara viridula 
nymphs (Heteroptera: Pentatomidae) with observations on its immature stages 
and development 131 

SOARES, L. A., L. R. S. ZANETTE, H. R. PIMENTA, A. GONgALVES, and R. P MARTINS. 

Nesting biology of Isodontia costipennis (Spinola) (Hymenoptera: Sphecidae) 245 

ZAMA, U., J. LINO-NETO, and H. DOLDER. Ultrastructure of spermatozoa in Plebeia 

(Plebeia) dron/nna Friese (Hymenoptera: Apidae: Meliponina) 261 



NOTE 



HUBER, J. T. and Z. BOUCEK. Pohjnema Haliday, 1833 (Insecta, Hymenoptera): 

designation of Polynema flavipes Walker, 1846, as the type species 280 



563 




Journal of 

Hymenoptera 
Research 



Volume 11, Number 1 April 2002 

ISSN #1070-9428 
CONTENTS 

EHLER, L. E., L. B. MALLER, and Y. HIROSE. Reproductive biology of Gnjou obeswii 

Masner (Hymenoptera: Scelionidae) 1 

ENGEL, M. S. The fossil pelecinid Pekcinopteron tiibiiUfoniic Brues in Baltic amber (Hyme- 
noptera: Pelecinidae) 5 

FINNAMORE, A. T Revision of the world genera of tribe Stigmini (Hymenoptera: Apoi- 

dea: Crabronidae: Pemphredoninae), Part 2. Species of IncastigDius Finnamore . . 12 

MANLEY, D. G. and J. P. PITTS. A key to genera and subgenera of Mutillidae (Hymenop- 
tera) in America north of Mexico with description of a new genus 72 

MATTHEWS, R. W. and I. D. NAUMANN. Descriptions and biology of nine new species 
of ArpmctopJiihis (Hymenoptera: Crabronidae), with a key to described Australian 
species 101 

SCHMIDT, S., D. R. SMITH, and J. MACDONALD. Review of the Australian subfamily 

Pteryperginae (Hymenoptera: Symphyta: Pergidae) 134 

SMITH, D. R., A. D. WRIGHT, A. WINOTAI, and R. DESMIER de CHENON. Studies on 
Neostromboceros albicomiis (Konow) (Hymenoptera: Tenthredinidae), a potential 
biological control agent for the Old World climbing fern, with notes on two other 
species of Neostromboceros 142 

WHITFIELD, J. B., A. BENZING, and R PONCE. Review of the Gh/ptapautelcs species 
(Hymenoptera: Braconidae, Microgastrinae) attacking noctuids in field crops in 
the Neotropical Region, with descriptions of two new species from the Ecuadorian 
Andes 152 



M 2 7 2002 

//SRARIFS 



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This issue was mailed 18 April 2002 



J. HYM. RES. 
Vol. 11(1), 2002, pp. 1-4 



Reproductive Biology of Gryon obesiim Masner 
(Hymenoptera: Scelionidae) 

L. E. Ehler, L. B. Maller, and Y. Hirose' 

Department of Entomology, University of California, One Shields Avenue, 
Davis, California 95616-8584, USA 



Abstract. — The reproductive biology of Giyou obcsiini Masner (Scelionidae) was investigated in 
the laboratory using eggs of Eusdiistits coiispcrsiis Uhler (Pentatomidae) as hosts. Females typically 
began ovipositing on the day of emergence and continued to oviposit for a mean of 40.7 days. 
Mean lifetime fecundity was 267 eggs/female. Mean daily fecundity was highest (—22 eggs/ 
female) on the first day. By days 10 and 17, a mean of ~50"'/() and ~757o of the eggs had been 
deposited, respectively. Sex ratio of progeny was female biased during the first part of the ovi- 
positional period and male biased during the latter part. The post-ovipositional period was rela- 
tively short (mean of 5.6 days). Mean female longevity was 47.3 days when males and hosts were 
present, compared to 61.7 days for females that were deprived of males and host eggs. Because 
G. obcsiini has a shorter generation time and a greater lifetime fecundity than £. conspersiis, it has 
great potential in augmentative biological control of this pest in crops such as processing tomato 
in northern California. 



Gn/on obcsii}}! Masner is a New World 
scelionid that parasitizes the eggs of stink 
bugs (Pentatomidae) (Johnson 1992). In 
the United States, it occurs primarily in 
southern areas, from Florida to California, 
where it is typically associated with hosts 
in the genera Euschistiis and TIn/aiita (Mas- 
ner 1983). In northern California, G. obes- 
uni is commonly reared from eggs of £. 
coiispcrsus Uhler (consperse stink bug) and 
T. pnlUdovirois (Stal) (red-shouldered stink 
bug) (Ehler 2000). However, its life history 
is poorly known, hi view of this, we ini- 
tiated laboratory investigations on G. obcs- 
iini, with particular emphasis on repro- 
ductive biology and patch-use patterns. 
The present paper is restricted to repro- 
ductive biology, and its relevance to the 
use of G. obcsiini in augmentative biologi- 
cal control of E. coiispcrsus on tomato in 
northern California. 



' Present address: 349 Asano, Munakata 811-3415, 
Japan (E-mail: yhirose@h7.dion.ne.jp) 



MATERIALS AND METHODS 

Laboratory cultures of £. coiispcrsus and 
G. obcsuni were established from field-col- 
lected material from the immediate vicin- 
ity of Davis, CA (Yolo County). Adults of 
E. coiispcrsus were collected from weedy 
hosts, and from cultivated crops such as 
tomato and dried bean. Male/female pairs 
were placed in individual 450-mI ice 
cream containers, supplied with fresh 
green-bean pods (Phascolus viilgnris L.) 
and raw sunflower seeds {Hcliantliiis iiii- 
iiiiiis L.), and held in a rearing room at 
25°C and 16:8 (L:D) photoperiod. Contain- 
ers were lined with paper towel which 
served as an ovipositional substrate. Con- 
tainers were inspected daily; newly de- 
posited eggs were cut out of the paper 
towel and fresh food was added as re- 
cjuired. Egg masses were stored at 10°C for 
later use in assessing fecundity of G. obcs- 
uni (usually <3 days). The culture of G. 
obcsuiu was established from individuals 
reared from eggs of southern green stink 



Journal of Hymenoptera Research 




Days After Emergence 



Fig. 1. Mean (SEM) dailv fecundity of G. obesuui. 



bug, Ncznm viridiila (L.). This culture was 
maintained continuously on eggs of E. 
couspcrsus in a rearing room at 25'C and 
10:14 (L:D) photoperiod. 

Newly emerged male/female pairs (n = 
10) of G. obesum were confined to 450-ml 
ice cream containers. A plastic petri dish 
cover was placed oxer the top of the con- 
tainer; fresh honev was streaked across 
the inside of the lid. Dead males were re- 
placed as needed with males of unknown 
age from the colony. Host egg masses 
were glued to a strip of paper that was 
inserted through an opening in the side of 
the container. Egg masses were changed 
at 24 h intervals. Individual females were 
initially exposed to 3-4 host egg masses 
(—40 eggs) per day; this was reduced to a 
single egg mass ( — 14 eggs) when daily fe- 
cundity declined to <10 eggs/female. Fol- 
lowing exposure, egg masses were re- 
moved, placed in glass vials with cotton 
plugs, and held in the rearing room for 
parasitoid emergence. Daily oviposition 
rate was based on the total number of 
eggs parasitized, including those from 
which adults did not emerge. Sex ratio 
was determined for egg clutches in which 
all progeny (emerged and not emerged) 
could be accurately sexed. Voucher spec- 
imens of G. olx'suni are deposited in the 



Bohart Museum of Entomology at the 
Uni\'ersitv of California, Davis. 

RESULTS AND DISCUSSION 

The daily production of progeny by G. 
obcsiiDi is summarized in Fig. 1. Nine fe- 
males oviposited on day one, and the re- 
maining female commenced ovipositing 
on day two. Mean (SEM) daily fecundity 
was highest on day one (21.9 ± 3.8 eggs/ 
female), ranged from 9.5 ± 1.8 to 15.5 ± 
4.3 eggs/female from day two through 
day 15, and gradually declined thereafter. 
Mean (SEM) lifetime fecunditv was 267 ± 
17.3 eggs/female. Of this total, -25% 
were deposited by day 5, —50% by day 10, 
and 75% by day 17. Mean (SEM) oviposi- 
tional and post-ovipositional periods were 
40.7 ± 3.2 and 5.6 ±1.9 days, respecti\'ely. 
Mean (SEM) longe\ity of females was 47.3 
±1.9 days (n = 7) compared to 61.7 ± 6.4 
days (n = 9) for females that were held 
under similar physical conditions but de- 
pri\'ed of males and host eggs (t = 1.914, 
df - 14, P = 0.076). 

Sex ratio of progeny was directly related 
to the age of the female (Fig. 2). It was 
strongly female-biased during the early 
stage of the oxipositional period, but grad- 
ually shifted to male-biased by the end of 
the ovipositional period. In the latter case. 



Volume 11, Number 1, 2002 




Days After Female Emergence 

Fig. 2. Relntionship between secondary sex ratio 
(proportion males) of progeny and age of female in 
G. obcsuiii. Regression/correlation statistics as fol- 
lows: Y = -0.05 + 0.026X, r- = 0.75, P = 0.0001. Only 
egg clutches in which all individuals could be accu- 
rately sexed are included; some points represent 
means oi two or more clutches. 



egg clutches were typically <5 eggs/ fe- 
male. This shift in sex ratio may have re- 
sulted from sperm depletion. However, as 
it was necessary to use males of unknown 
age in some cases, this hypothesis cannot 
be adequately tested here. 

Waage (1982) noted that maximum life- 
time fecundity of scelionids probably 
ranges from 50-150 eggs/female. Thus, 
lifetime fecundity for G. obcsiiiu (>250 
eggs /female) would appear to be excep- 
tionally high for a scelionid. It also is 
much greater than that of two Nearctic 
scelionids that commonly parasitize £. 
conspcrsiis in northern California— i.e., 
—40 to —80 eggs/ female for Tckvionius 
pod i si Ashmead (Yeargan 1982, Orr and 
Boethel 1990) and -65 for Trissolcus cits- 
chisti (Ashmead) (Yeargan 1982). The 
same holds for Cryoi pcinisi/lvniiicuDi 
(Ashmead), a Nearctic egg parasite of 
squash bug, Aunsn tristis DeGeer — i.e., 
-80 eggs/female (Nechols ct nl. 1989). 
Also, niean ovipositional period (40.7 
days) and female longevity (47.3 days) for 
G. obcsiim are greater than for T. podisi 
( — 8 and —12 days, respectix'ely) (Yeargan 



1982, Orr and Boethel 1990), T. ciischisti 
(—28 and —35 days, respectively) (Year- 
gan 1982), and G. pciuisylviniiciun (22 and 
-40 days, respectively) (Nechols ct al. 
1989). 

Based on reproductive biology of the 
three scelionid parasitoids of E. coiispersiis 
in northern California, G. obcsuni would be 
the clear choice for augmentative release 
against this pest in tomato. It has the high- 
est lifetime fecundity, longest oviposition- 
al period, and greatest female longevity of 
the three parasitoids, and thus could be 
expected to have the greatest impact on 
pest density. Also, G. obcsuni has a repro- 
ductive advantage over that of the pest. 
Lifetime fecundity of £. conspwrsus is —225 
eggs /female at 27°C (Hunter and Leigh 
1965, Toscano and Stern 1976), compared 
to >250 for G. obcsuni. Also, generation 
time for G. obcsuni is much shorter — i.e., 
— 15 days (unpublished data) compared 
to —50 tc^ —55 ciays for E. coiispcrsus 
(Hunter and Leigh 1965, Toscano and 
Stern 1976). Thus G. obcsuin could com- 
plete at least three generations to each 
host generation. Finally, it could be ex- 
pected to deposit —50% of its eggs in the 
first 10 days after emergence, further en- 
hancing its impact in augmentative bio- 
logical control. 

ACKNOWLEDGMENTS 

We thank M. J. Tauber and C. A. Tauber of Cornell 
University for critical review of the manuscript. This 
research was supported by Regional Research Project 
W-185. 

LITERATURE CITED 

Ehler, L. E. 2000. Fnnuscapc Ecology of Stink Bu;^> in 
Noriiiern California. Memoirs, Thomas Say Pub- 
lications in Entomology. Entomological Society 
of America, Lanham, MD. 59 pp. 

Hunter, R. E. and T. F. Leigh. 1965. A laboratory life 
history of the consperse stink bug, Eiischigtit^ con- 
spcrsiis (Hemiptera: Pentatomidae). Annah of the 
Rntoniolo^ical Society of America 58; 648-649. 

Johnson, N. F. 1992. Catalog of World Species of Proc- 
totrupoidea. Exclusive of Playtgastridae (Hyme- 
noptera). Memoirs, American Entomoloj^ical Insti- 
tute, No. 51. Gainesville, FL. 



Journal of Hymenoptera Research 



Masner, L. 1983. A revision of Gri/oii Haliday in 
North America (Hymenoptera: Proctotrupoidea: 
Scelionidae). Tlic Cninniiaii E)itoiiiolo;,^ist 115: 123- 
174. 

Nechols, ]. R., J. L. Tracy and E. A. Vogt. 1989. Com- 
parative ecological studies of indigenous egg 
parasitoids (Hymenoptera: Scelionidae; Encyrti- 
dae) of the squash bug, Aiiasn tristis (Hemiptera: 
Coreidae). journal of the Ktiusns Eiitoi}iolo;^icnI So- 
cii'ti/ 62: 177-188. 

Orr, D. B. and D. ]. Boethel. 1990. Reproductive po- 
tential of TclcJioiuus cristntiis and T. podisi (Hy- 
menoptera: Scelionidae), two egg parasitoids of 



pentatomids (Heteroptera). Annals of the Ento- 
mological Society of America 83: 902-905. 

Toscano, N. C. and V. M. Stern. 1976. Development 
and reproduction of Euschistus co)ispcrsiis at dif- 
ferent temperatures. Annals of the Entomoh\^ical 
Society of America 69: 839-840. 

Waage, J. K. 1982. Sib-mating and sex ratio strategies 
in scelionid wasps. Ecolo;^ical Entomolo;;i/ 7: 103- 
112. 

Yeargan, K. V. 1982. Reproductive capability and lon- 
gevity of the parasitic wasps Telenoiinis podisi 
and Trissolcus eiischisti. Annah of the Entomological 
Society of America 75: 181-183. 



J. HYM. RES. 
Vol. 11(1), 2002, pp. 5-11 

The Fossil Pelecinid Pelecinopterou tubiiliforme Brues in Baltic 
Amber (Hymenoptera: Pelecinidae) 

Michael S. Engel 

Department of Ecology and Evolutionary Biology, and Division of Entomology, Natural History 

Museum and Biodiversity Research Center, Snow Hall, 1460 Jayhawk Boulevard, University of 

Kansas, Lawrence, Kansas 66045-7523, USA 



Abstract. — The fossil pelecinid Pckxiiioptcnvi tiibiilifoniic Brues (Proctotrupoidea) is redescribed 
and figured based on a single, complete male preserved in middle Eocene (Lutetian) Baltic amber. 
Brues' original material is missing but comparison with his description and figures allows for 
positive identification of the species. Since the type material for this genus and species are un- 
traceable and presumably destroyed the new specimen is herein designated as a neotype for the 
purpose of stabilizing the nomenclature and identity of this, the only definitive fossil of the family 
Pelecinidae. 



The family Pelecinidae comprises the gi- 
ants of the superfaniiiy Proctotrupoidea 
with slow-flying indiviciuals ranging in 
size from 25 to 60 mm in total length. The 
family is today represented by only three 
extant species, all restricteei to the Western 
Hemisphere — Pclcciuiis pohitiirator (Drury) 
is known from southeastern Canada, the 
eastern United States and Mexico, and 
south to northern Argentina; P. dichroiis 
Perty in southeastern Brazil, Paraguay, 
Uruguay, and northern Argentina; and P. 
tliorncicus Klug presently known only 
from western Mexico. Little is known of 
Peleciiius biology aside from some melo- 
lonthine host records for P. pwh/turator. In- 
dividuals of P. pwlyturntor have been 
reared from larvae of several Pliyllophagn 
species (Coleoptera: Scarabaeidae) (see 
summary in Johnson and Musetti 1999). 
The three species were described and a 
key presented for their identification by 
Johnson and Musetti (1999). 

Johnson (1998) recently reviewed the 
two fossil species for the family: Pclccinop- 
tcrcvi tiibiilifoniie Brues (1933) in Baltic am- 
ber and Iscopiiiiis bnissicus Kozlov (1974) 
preserved as a compression fossil from the 



Lower Cretaceous of the Transbaikal re- 
gion. Based on considerable differences in 
wing venation and uncertain affinity to 
Peleciiius or other proctotrupoids, Johnson 
(1998) rightfully removed Iscopinus from 
the Pelecinidae s.str. and considered it as 
a family of indeterminate position within 
the Proctotrupoidea. This action left the 
Eocene genus Pelecinopterou as the sole 
fossil representative for the Pelecinidae. 
Unfortunately, the two males and single 
female upon which Brues (1933) based his 
original description were from the ill-fated 
collections of the Albertus Universitat in 
Konigsberg (today Kaliningrad, Russia). 
During World War II this collection was 
destroyed by fire. St^me portion of the col- 
lection was spared and today resides in 
the Institut und Museum f'tir Geologic 
und Palaontologie, Gottingen. A personal 
investigation of this collection was made 
in July of 1999 but no material of Peleci- 
uopterou could be discovered (other Euro- 
pean institutions with amber collections 
were visited at the same time and addi- 
tional Konigsberg material was not locat- 
ed). 

Herein 1 provide a new description and 



Journal of Hymenoptera Research 








\ \ 



'k 




I ie,. I. niotomicrogmph of Prlrcinopteroii lubulifonuc Bruos (AMNH). 



Volume 11, Number 1, 2002 



figures for a complete male of P. tulmUfor- 
mc recently identified in middle Eocene 
Baltic aniber and designate this specimen 
as a neotype for the species. Forniat for the 
description generally follows that em- 
ployed by Johnson and Musetti (1999) for 
living pelecinids so as to aid comparison 
with Pdccinus. Measurements were made 
using an ocular micrometer c^n an Olym- 
pus SZX12 stereomicroscope and should 
be cc^nsidered somewhat approximate 
since the c^ptimal angle for some metrics 
was not always achievable. Microphoto- 
graphs were prepared using a Microptics 
ML-1000 digital imaging system. The age 
and origin of Baltic amber has been re- 
cently reviewed in Engel (2001). 

SYSTEMATIC PALEONTOLOGY 

Genus Pelecinopterou Brues 

Pelecinopteron Brues 1933: 19. Type species: Pe- 
lecinopteron tiibulifonuc Brues 1933, monoba- 
sic and original designation. 

Diagnosis. — Male. Inner margins of 
compound eyes very slightly convergent 
below, essentially parallel; maxillary pal- 
pus 5-segmented; labial palpus 3-seg- 
mented [I could not discern a fourth, 
short, basal segment alluded to by Brues 
(1933)]; mandible bidentate, teeth short 
and equal in length, outer surface without 
dense, elongate setae; niandibles broadly 
overlapping. Clypeus convex, with coarse, 
faint punctures scattered over surface (dis- 
tinctly not strongly punctured), without 
elongate setae, apical margin relatively 
straight. Ocelli positioned in equilateral 
triangle near top of vertex, median ocellus 
at upper tangent of compound eyes. Oc- 
cipital carina strong, distinctly present 
both medially and laterally. Antenna fili- 
form, 13-segmented; positioned slightly 
below midpoint of face, separated from 
base of clypeus by ca. 1.75X antennal 
socket diameter; combined lengths of 
scape and pedicel much shorter than first 
flagellomere, tlagellomeres elongate, basal 
four flagellomeres with length ca. 4.5- 



6.5 X width, following four segments with 
length ca. 4x width, distal three segments 
with length ca. thrice width. Pronotum an- 
nular, dorsally with posterior section trap- 
ezoidal, this section anteriorly bordered 
by strong, transverse carina; anterior to ca- 
rina pronotum gently sloping down to 
short anterior collar. Notauli formed of 
posteriorly converging, strong, crenulate 
impressions, confluent posteriorly; mesos- 
cutum and scutellum separated by narrow 
suture, suture bordered by row of large, 
strong foveae on scutellum; axillae nar- 
row; scutellum weakly arched; metano- 
tum short. Mesepisternum with transverse 
furrow extending from faint episternal 
groove posteriorly, not reaching mes-me- 
tepisternal suture. Propodeum elongate; 
strongly and coarsely sculptured; sparsely 
setose. Tibial spur forn^iula 1-2-2; metati- 
bia gently expanded apically, metabasitar- 
sus distinctly elongate, longer than three 
immediately following tarsal segments 
(i.e., length of tarsal segment 1 ~ com- 
bined lengths of tarsal segnients 2, 3, and 
4); second tarsomere one-half length of 
metabasitarsus; fourth tarsomere extreme- 
ly short, with inner apical margin project- 
ing underneath fifth tarsal segment. Fore- 
wing with only two tubular veins (C and 
Sc4-R); pterostigma elongate, tapering to 
point on anterior wing margin; R not ex- 
tending beyond pterostigma; first abscissa 
of Rs slightly angled toward wing base, 
subequal in length to basal vein (i.e., first 
free abscissa of M); r-rs arising slightly ba- 
sad pterostigmal midpoint (distad pteros- 
tigmal midpoint in Pdccinus); Rs forking 
slightly basad pterostigmal apex and dis- 
tad forewing midpoint, forming two 
branches, Rsl and Rs2, each branch equal- 
ly pigmented and reaching to wing apex; 
Rsl arching anteriorly before extending to 
wing apex; medial cell elongate; Cu reach- 
ing wing apex, slightly more heavily pig- 
mented near wing margin than distalmost 
abscissae of Rsl, Rs2, and M; 2cu-a slight- 
ly distad Im-cu; veins more strongly pig- 
mented in basal two-thirds of wing (i.e.. 



Journal of Hymenoptera Research 



from slightly beyond pterostigmal apex to 
base of wing) except around second ab- 
scissa of Rs + M and anal vein with asso- 
ciated crossveins (i.e., Icu-a and 2cu-a) 
very faintly indicated; membrane hyaline; 
venational details presented in figure 4. 
Hind wing without venation except C 
along anterior margin; without closed 
cells; membrane hyaline. Metasoma elon- 
gate; sixth metasomal segment swollen 
and enlarged, with strong teeth along lon- 
gitudinal n^iidline of sternuni, first tooth at 
midpoint of sternal length, second tooth 
near apical fourth; sixth tergum and sixth 
sternum partially fused (i.e., suture be- 
tween them exceedingly faint), same for 



seventh metasomal segment; seventh me- 
tasomal segment generally falcate; para- 
meres elongate. Female. Surviving speci- 
mens unknown; based on descriptive de- 
tails in Brues' (1933) original description 
the female is generally as described for the 
male herein except for sexual differences. 
This will require confirmation when new 
material of the female sex is ciiscovered. 

Coinuicuis. — Brues (1933) originally pro- 
posed a separate family, Pelecinopteridae, 
for this genus while noting its strong af- 
finity to Pelecinidae. Owing to the enor- 
mous similarity of PcIcciiiopHcnvi with Pt'- 
leciiiiis I agree with most authors (e.g., 
Johnson 1998) that the two genera shc^uld 
be placed in a single family. 



Key to Genera of Pelecinidae 

1. Antenna 14-segmented; malar space well-developed; r-rs arising in distal half of pterostig- 
ma; forewing with infuscation (particularly in costal cell, along anterior margin, and at 
apex); metabasitarsus distinctly shorter than following tarsomere; male metasoma clavate; 
body size large, ca. 25-60 mm (extant; Western Hemisphere) Pelecinits Latreille 

- Antenna 13-segmented; malar space extremely short; r-rs arising in basal half of 

pterostigma; forewing apparently without infuscation; metabasitarsus distinctly longer 
than following tarsomere; male metasoma elongate; body size moderate, ca. fO-15 mm 
(early Cenozoic amber; Europe, northern Asia) Pclccinopteron Brues 



Pelecinoptcron tuhnliforme Brues 

(Figs. 1-4) 

Pelecinoptcron tiibiiliforme Brues 1933: 20. Kozlov 
1974: 145 (translated version 1974: 137) [par- 
tial Paleocene amber specimen]. Johnson 
1998: 2 [descriptive notes]. 

Diagnosis. — As for the genus (see 
above). 

Descriptioi. — As described for the genus 
with the following additions: Male. Total 
body length (excluding antennae) 10.9 
mm; forewing length 4.3 mm; head length 
1.5 mm (head width indeterminate owing 
to angle from which front of face can be 
seen through amber surface); malar space 
length 0.07 mm; length of compound eye 
1.1 mm; metasomal length 8.4 mm; length 
of first metasomal segment 1.0 mm; length 



of second metasomal segment 1.0 mm; 
length of third metasomal segment 1.0 
mm; length of fourth metasomal segment 
0.8 nim; length of fifth metasomal segment 
1.3 mm; length of sixth metasomal seg- 
ment 1.9 mm; length of seventh metaso- 
mal segment 1.4 mm. Coloration of integ- 
ument not well preserved, where evident 
apparently dark brown to black through- 
out. 

Clypeus with sparse, faint, rather small, 
coarse punctures, integument otherwise 
sniooth; remainder of head with minute, 
scattered punctures, integun"ient between 
smooth. Dorsal-facing surface of proncv 
tum with posterior portion separated from 
collar by transverse carina, integument of 
dorsal surface with minute punctures sep- 
arated by a puncture width, integument 



VOLUMH 11, NUMBFR 1, 2002 








,* 




\ 



•#^ 




X. 




Figs. 2-3. Photomicrographs of Pclcci)Joptcnvi tubulifonuc Brucs (/\M,\H). 2. Lateral view of mesosoma and 
head. 3. Lateral \'iew of distal metasomal segments (from left to right = segments 7, 6, 5, 4, and apex of 3). 



10 



Journal of Hymenoptera Research 




Fig. 4. Forewing \'enation of Pelccuioptcnvi tiibulifoiuic Brues (AMNH). Scale bar = 0.5 mm. Dashed line 
indicates an alar fenestra. 



between smooth or slightly imbricate; lat- 
eral surface of pronotum with dorsal third 
sniooth except along border with mesos- 
cutum with niinute punctures separated 
by a puncture width or slightly more, in- 
tegument below smooth patch pronotum 
with large and strong punctures, punc- 
tures nearly confluent, integument be- 
tween punctures (where evident) smooth. 
Mesoscutum with crenulations along lat- 
eral and anterior borders; notauli strongly 
impressed and crenulate, fusing posteri- 
orly; integument in crenulations smooth, 
otherwise integument with n^iinute punc- 
tures separated by a puncture width or 
slightly n^iore; tegula with minute punc- 
tures separated by a puncture width, in- 
tegument between punctures smooth; scu- 
tellum with strong and deep foveae along 
margins to form a small, medial, horizon- 
tal surface, integument in foveae smooth, 
on medial surface with minute punctures 
separated by a puncture width or slightly 
more, integument between punctures 
smooth. Mesepisternum with large and 
strong punctures, punctures nearly con- 
fluent, integument between punctures 
smooth; with transverse depression ante- 
riorly connecting to short and more faint 
episternal groove (posteriorly this groove 
does not reach to the suture between the 
mes- and metepisternum), integument in- 
side of groove smooth, above groove in- 
tegument with minute punctures separat- 



ed by a puncture width or less, integu- 
ment between punctures smooth. Metepis- 
ternum sculptured as on mesepisternum 
below transverse groove. Propodeum 
sculptured as on mesepisternum below 
transverse groove except posteriorly punc- 
tures fusing to form large areolae. Meta- 
soma with minute punctures widely scat- 
tered, integument otherwise smooth. Setae 
generally minute (less than a single ocellar 
diameter in length) and sparse except on 
legs, metasonia, and borders of pronotum 
and mesoscutum slightly more extensive 
but distinctly not dense. Female. Surviv- 
ing specimens unknown; froni the few de- 
scriptive notes provided by Brues (1933) 
females are generally as described for the 
male (above) aside from the typical sexual 
differences and perhaps some slight vari- 
ations in sculpturing of the mesosoma. 

Material — Neotype (here designated). 
Male, Baltic amber, middle Eocene (Lute- 
tian); labeled "Neotype, Pclcci)wptcron tub- 
ulifoniic Brues, desig. M. S. Engel [red la- 
bel]"; deposited in the Amber Collection 
of the Division of Invertebrate Zoology, 
American Museum of Natural History, 
New York. This specimen is designated in 
accordance with Article 75.3 (ICZN 1999) 
and for the express purpose of clarifying 
and stabilizing the taxonomic status of P. 
iubuUfornic. The new specimen originates 
from the same deposits as the original se- 
ries (i.e., the middle Eocene "Blue Earth" 



Volume 11, Number 1, 2002 



11 



deposits of northern Europe, from which 
Baltic amber originates: see Engel 2001). 
Althc^ugh the holotype was originally a fe- 
male, two males were also described and 
the current specimen corresponds in all 
observable details to those features de- 
scribed for the male by Brues (1933). Thus, 
in accordance with Article 75.3.5 the neo- 
type may be based on a different sex (in 
this instance, a male). 

Coifuiiciits. — As mentioned above, the 
original material upon which Brues (1933) 
based his description is missing (and per- 
haps destroyed with the bulk of the Ko- 
nigsberg collection during World War 11). 
A partial specimen consisting of only a 
male metasoma is known in Paleocene 
amber from Sakhalin, Russia (Kozlov 
1974, Johnson 1998). No other specimens 
are presently recorded for this taxon. 
Thus, the specimen described herein is the 
only complete, surviving individual for 
the species and the only one originating 
from the same deposits as the original 
type series. When a more complete speci- 
men of the species from Paleocene amber 
of Russia is discovered it may prove to be 
a separate species from P. tubiilifonne. For 
example, the photograph published by 
Johnson (1998) does not clearly show the 
ventral teeth on the swollen sixth meta- 
somal segment. Other differences may 
come to light with more completely pre- 
served material. 



ACKNOWLEDGMENTS 

I am grnteful to Hans Jahnke for proxiding rooms 
in the Institut und Museum fur Geologie und Palaon- 
tologie, for hosting my visit to Gottingen in 1999, and 
for genera! information on the amber collection; to 
David A. Grimaldi for bringing this specimen to my 
attention; to Tam C. Nguyen with help accessing lit- 
erature; and to Allan H. Smith-Pardo, E. Eric Grissell, 
and two anonymous reviewers for commenting on 
the manuscript. This is contribution Nr. 3282 of the 
Division of Entomology, Natural History Museum 
and Biodiversity Research Center, University of Kan- 



LITERATURE CITED 

Brues, C. T. 1933. The parasitic Hymenoptera of the 
Baltic amber: Part 1. Bcnistciii-Forschiui^^cii 3: 4- 
178. 

Engel, M. S. 2001. A monograph of the Baltic amber 
bees and evolution of the Apoidea (Hymenop- 
tera). Bulletin of the Aiucrican Miisciiiii of Natural 
Histon/ 259: 1-192. 

International Commission on Zoological Nomencla- 
ture. 1999. International Code of Zoological Nonien- 
elature [4"' Edition]. International Trust for Zoo- 
logical Nomenclature; London, United Kingdom; 
xxix + 306 pp. 

Johnson, N. F. 1998. The fossil pelecinids Pelecinopter- 
on Brues and Iseopinus Kozlov (Hymenoptera: 
Proctotrupoidea: Pelecinidae). Proceedi}igs of the 
E)itoiuological Soeieti/ of Washington 100(1): 1-6. 

Johnson, N. F., and L. Musetti. 1999. Revision of the 
proctotrupoid genus Peleei)uis Latreille (Hyme- 
noptera: Pelecinidae). journal of Natural History 
33: 1513-1543. 

Kozlov, M. A. 1974. [An early Cretaceous ichneumon 
of the family Pelecinidae (Hymenoptera, Peleci- 
noidea)]. Paleo)itologicheskii/ Zhurnal 1974(1): 144- 
146. [In Russian: English translation in Paleonto- 
logieal journal 8(1): 136-138] 



J. HYM. RES. 
Vol. 11(1), 2002, pp. 12-71 

Revision of the World Genera of Tribe Stigmini (Hymenoptera: 

Apoidea: Crabronidae: Pemphredoninae), Part 2. Species of 

Incastigmtis Finnamore 

Albert T. Finnamore 

Invertebrate Zoology, Provincial Museum of Alberta, 12845-102 Avenue, 
Edmonton, Alberta, Canada T5N 0M6, E-mail: Albert. Fimnamore@gov.ab.ca 



Abstract. — liicnsti^nius is a Neotropical genus previously known from 2 species. In this paper 
21 species of bicnsti^iiiiis are described as new: aylaxiter (Argentina, Bolivia, Brazil), caelukhiis 
(Peru), cearaciisis (Brazil), ceromtis (Peru), cJiiiicha (Ecuador, Venezuela), ictericornis (Argentina, 
Paraguay, Bolivia, Brazil, Peru, Ecuador), igititlwrnx (Costa Rica, Panama), ipliis (Brazil, Paraguay, 
Bolivia, Peru), kiinkopteryx (Brazil, Bolivia, Peru, Ecuador, Colombia), mauracis (Brazil, Bolivia, 
Peru, Ecuador), m\/staxalbus (Mexico, Guatemala, Honduras, El Salvador, Costa Rica), tuytior 
(Brazil, Bolivia, Ecuador, Colombia, Venezuela), paranicus (Argentina, Bolivia, Brazil), propho- 
rodontis (Brazil, Bolivia, Ecuador, Colombia, Panama, Venezuela, Trinidad and Tobago), pyciiog- 
lypticiis (Brazil), pyrrhopyxis (Peru, Ecuador, Colombia, Costa Rica, Trinidad and Tobago), strcp- 
siliiicntiis (Venezuela), simiccrns (Brazil), trichodoceriis (Brazil, Paraguay, Peru, Ecuador, Colom- 
bia, Venezuela, Suriname, Trinidad anci Tobago), tirqicus (Brazil) and zephyriis (Mexico, Guate- 
mala, Nicaragua, Costa Rica, Panama). Stiginus hcxagoiinlis Fox (Colombia, Ecuador, Peru, Brazil) 
and S. neotropiais Kohl (Texas to Argentina) new combinations, are transferred to liicnstigiiius. 
Additionally, a key is provided to all species. 



The genus hicastiguiiis, with 25 currently 
recognized species, was ciescribeci to de- 
fine a lineage of Neotropical Stigmini 
based on a phylogenetic analysis of the 
world taxa (Finnamore 1995). Its speci- 
mens are the most common Neotropical 
Stigmini in museum collections. The ge- 
nus ranges from southern Texas to Argen- 
tina, but is not known north of the Lesser 
Antilles in the Carribean, or from Chile. 
Nothing is knc^wn of the biology and be- 
haviour of Incastigmus, but it is likely that 
all species construct nests in twigs and 
provision with aphids, as do most species 
in related genera. Several species in relat- 
ed genera nest in pre-existing cavities and 
some morphological and anecdotal evi- 
dence suggests Pdnistigimis species may be 
sand-nesting (Finnamore 1995). The title 
of the present paper reflects recent chang- 
es to the classification of apoid wasp lin- 
eages which, among other things, placed 



the Pemphredoninae in the family Cra- 
bronidae (Melo 1999). 

METHODS 

Terminology generally follows Bohart 
and Menke (1976), but in some cases 
needs clarification. Morphological terms 
are listed belc^w: 

Apprcsscii sctnc: setae forming an angle 
close to 0" with the body surface. 

Lateral Sphere of propodeiini: convex area 
c^f propodeum between propodeal enclo- 
sure and side (Gittins 1969). 

LOD: maximum diameter of lateral 
ocellus. 

Mesosoiiia: the thorax plus the propo- 
deum. 

Metasoiiia: the apparent abdomen con- 
sisting of the abdomen excluding the first 
segment or propocieum. 

Mieropore field: a grouping of minute 
pores usually visible only with scanning 



Volume 11, Number 1, 2002 



13 



electron microscope, but by stereomicro- 
scope with diffusing filter apparent as a 
discrete microsculpture patch or line on 
upper frons, usually between the lateral 
ocellus and the eye. 

Microsculpture: minute sculpture im- 
parting a dull appearance to the body. 

OOD: ocellocular distance, the least dis- 
tance between lateral ocellus and eye. 

Prcoiuaular area: area of mesopleuron an- 
terior to omaulus ( = preomaulal area of 
Bohart and Menke (1976)). 

Transverse groove: on pronotal dorsum, 
the transverse groove immediately poste- 
rior to the transverse carina. 

Descriptions of all included species are 
provided based on the material examined. 
In species demonstrating variability, de- 
scriptions are based on representatives of 
the most prevalent phenotype with varia- 
tion noted throughout the description. 
Collection data for the holotypes are pre- 
sented as they appear on the label; thus 
several spellings for the same locality and 
collectors, and several formats for date of 
collection may be encountered. Collection 
data for paratypes are presented in a stan- 
dard format of descending political units. 
Square brackets [ ] are used to indicate 
misspelling of localities, inability to con- 
firm placement of a locality within a po- 
litical unit, or when two or more localities 
with identical spelling exist within a po- 
litical unit and the label data was insuffi- 
cient to indicate which was intended (e.g., 
PERU: Ucayali: [San Pedro] — any of 4 lo- 
calities named San Pedro within Ucayali 
Department, 47 localities within Peru). For 
previously described species only the col- 
lection localities and museums are listed. 

The 2,043 specimens examined during 
this study were obtained from the follow- 
ing 31 institutions (the abbreviation pre- 
ceding the institution is that used in the 
text to designate repositories): 

AEIC — American Entomological Insti- 
tute, Gainesville, Florida, U.S.A. (D. 
Wahl). 

ANIC — Australian National Insect Col- 



lection, CSIRO, Canberra, A.C.T., Austra- 
lia. (l.D. Naumann). 

BMNH— The Natural History Museum, 
London, United Kingdom. (C. Taylor, C.R. 
Vardy). 

BPBM — Bernice P. Bishop Museum, 
Honolulu, Hawaii, U.S.A. (G.M. Nishida). 

CASC — California Academy of Scienc- 
es, Golden Gate Park, San Francisco, Cal- 
ifornia, U.S.A. (W.J. Pulawski). 

CMNH — Carnegie Museum of Natural 
History, Pittsburg, Pennsylvania, U.S.A. 
(J.E. Rawlings). 

CNCl — Canadian National Collection, 
Agriculture and Agri-Food Canada, Otta- 
wa, Ontario, Canada. (J. Huber, L. Mas- 
ner). 

CSUC — Department of Entomology, 
Colorado State University, Fort Collins, 
Colorado, U.S.A. (H.E. Evans). 

FSAG — Collections Zoologiques, Facul- 
te des Sciences Agronomiques, Gembloux, 
Belgium. (J. Leclercq). 

FSCA — Florida State Collection of Ar- 
thropods, Gainesville, Florida, U.S.A. (L. 
Stange, J. Wiley). 

HNHM — Zoological Department, Hun- 
garian Natural History Museum, Buda- 
pest, Hungary. (J. Papp). 

IIES — Instituto de Investigaciones En- 
tomologicas Salta 'TNESALT", Salta, Ar- 
gentina. (M.A. Fritz). 

IMLA — Fundacion E Instituto Miguel 
Lillo, Universidad Nacional de Tucuman, 
Tucuman, Argentina. (A. Willink). 

IZAV — Instituto de Zoologia Agricola, 
Universidaci Central de Venezuela, Mara- 
cay, Aragua, Venezuela. (J. Luis Garcia). 

LACM — Los Angeles County Museum 
of Natural History, Los Angeles, Califor- 
nia, USA. (R. Snelling). 

MACN — Museo Argentino de Ciencias 
Naturales, Buenos Aires, Argentina. (A. 
Roig Alsina). 

MCZC — Museum of Comparative Zo- 
ology, Harvard University, Cambridge, 
Massachusetts, U.S.A. (C. Vogt). 

MJMO — Decanato de Agronomia, Univ- 



14 



Journal of Hymenoptera Research 



ersidad Centre Occidental, Barquisimeto, 
Lara, Venezuela. (E. Yustiz). 

MPEG — Museu Paraense Emilio Goeldi, 
Belem, Para, Brazil. (W.L. Overal). 

MTEC — Entomology Collection, Mon- 
tana State University, Bozeman, Montana, 
U.S.A. (M. Ivie). 

MUCR — Museo de Insectos, Universi- 
dad de Costa Rica, Ciudad Universitaria, 
Costa Rica. (P. Hanson). 

NHMW — Naturhistorisches Museum 
Wien, Wien, Austria. (M. Fischer). 

OSUO — Department of Entomology 
Collection, Oregon State University, Cor- 
vallis, Oregon, U.S.A. (G.R. Ferguson). 

PMAE — Provincial Museum of Alberta, 
Edmonton, Alberta, Canada. 

RMNH — Nationaal Natuurhistorische 
Museum, Leiden, The Netherlands. (C. 
van Achterberg). 

SEMC — Snow Entomological Museum, 
University of Kansas, Lawrence, Kansas, 
U.S.A. (R.W. Brooks). 

SDMC— San Diego Natural History 
Museum, San Diego, California, U.S.A. 
(D.K. Faulkner). 

SMTD — Staatliches Museum fur Tier- 
kunde, Dresden, Germany. 

USNM— Smithsonian histitution, Wash- 
ington D.C., U.S.A. (K.V. Krombein, A.S. 
Menke). 

ZMAN — Instituut voor Taxonomische 
Zoologie, Universiteit van Amsterdam, 
Amsterdam, Netherlands. (W. Hogenes). 

ZMUM — Zoological Museum, Univer- 
sity of Moscow, Moscow, Russian Feder- 
ation. (A.V. Antropov). 

INCASTIGMUS Finnamore 

liiaistl^^Diiis Finnamore 1995:234. Type species: 
hicastigimis iiiti Finnamore 1995:235, by orig- 
inal designation. 

Rccogiiiticvi. — Specimens of bicasti^^imis 
can usually be recognized by the presence 
on the scutum of a median groove or pos- 
teromedian pit, a unique structure within 
western hemisphere Stigmini. The groove 
or pit must not be confused with the me- 



dian, or posteromedian ridge or multiple 
ridges in some species of Neotropical Sti^^- 
miis. The following combination will pro- 
vide assurance of generic assignnient for 
all known species: hind wing media di- 
verging before cu-a; acetabular carina pre- 
sent; mandibles tridentate in both sexes 
with apicoventral tooth acute in female; 
and scutum with a median groove or pos- 
teromedian pit. 

Since publication of Part 1 of this work 
(Finnamore 1995) I have found several 
males of Llnqlnistigmus iimiitiinti Finnamore 
which can be confused with those liiciistig- 
niiis in which the scutal groove is reduced. 
Male iiKiiitniiti sometimes have a series of 
short, evanescent, longitudinal striae 
along the posterior scutal margin in which 
the median pair mimic the posteromedian 
pit of some Iiicastigimis. The large bilobed 
labrum (lobes broad and separated by a 
slight emargination) in iiimitiiiiti indicates 
it is a Llaqhastigimis, whereas lucastigiuus 
possess a relativ^ely small quadrilobeci la- 
brum (median lobes narrow and separat- 
ed by a deep median notch). Male iiiiiiitnii- 
ti have been included in the following key 
to species of liicnstigiiius in order to avoid 
spreading mandibles to expose the labrum 
for generic determination. Female iiunitnn- 
ti possess a large bilobed labrum and a 
slightly enlarged, blunt, apicoventral man- 
dibular tooth, both diagnostic characters 
for Llnqlmstignius, and should not be con- 
fused with Inciistigiiiiis (apicoventral man- 
dibular tooth acute) in the key to genera 
of Stigmini (Finnamore 1995). In addition, 
female iimiitniiti lack any trace of a pos- 
teromedian scutal pit and the lateral clyp- 
eal teeth meet or exceed the median clyp- 
eal teeth in length. In liicastigiinis the lat- 
eral clypeal teeth are smaller than the me- 
dian teeth. 

Description. — Head: Labrum 4-lobed, 
with narrow deep median emargination; 
mandibular apex of both sexes tridentate; 
apicoventral mandibular tooth in female 
acute; inner basal mandibular tooth ab- 
sent; clypeal apex in male without bev- 



Volume 11, Number I, 2002 15 

elled modifications, in female with 4 teeth; omaulus continuous with acetabular cari- 

interantennal tubercle and frontal line ab- na; scrobal sulcus present; hypoepimeral 

sent; micropore field present between lat- area without coarse sculpture; mid basi- 

eral ocellus and eye; eye inner margins tarsus of male elongate, as long as next 3 

converging below; eyes at most partially tarsomeres combined; posterior margin of 

margined by a carina; occipital carina pre- hind tibia with 2 or 3 spines; fore wing 

sent, complete, not joining hypostomal ca- asetose in cellular area; hind wing media 

rina, simple in female, raised, foveolate in diverging before cu-a; hind wing subme- 

male. Mesosoma: Scutum with median dian cell of normal size, not reduced. Me- 

groove or at least posteromedian pit; no- tasoma: Petiole carinate; pygidial plate 

tauli often elongate, reaching posterior narrow, absent in male; digitus longer 

scutal margin; acetabular carina present; than cuspis, clubbed. 



KEY TO SPECIES OF INCASTIGMUS 

1 Male: antenna with 11 flagellomeres, metasoma with 7 exposed terga 2 

1' Female: antemia with 10 flagellonieres, metasoma with 6 exposed terga 29 

2 Vertex with micropore field present as a small oval or circular patch between lateral 
ocellus and eye margin (Fig. 14) 3 

2' Vertex with micropore field forming a narrow linear furrow between lateral ocellus and 

eye margin (Figs. 50, 58) 24 

3 Scutum with at most median scutal groove complete (Figs. 3, 15, 43, 71, 79); notauli 
attenuated posteriorly, not reaching posterior scutal margin except in a few cases by weak 
extensions; median scutal groove usually incomplete, sometimes reduced to a small pit 

on posterior scutal margin, or absent, or sometimes contiguous with admedian lines .... 4 
3' Scutum with 3 complete grooves (Figs. 23, 31); notauli reaching posterior scutal margin, 
not attenuated posteriorly; median scutal groove reaching admedian lines, not attenuated 
anteriorly 20 

4 Preomaular area sparsely setose with underlying sculpture clearly evident; median scutal 
groove present at least as a small pit on posterior margin; labrum quadrilobed 

lucastiginiis 5 

4' Preomaular area densely setose, with underlying sculpture obscured; posterior scutal 

margin with a series of evanescent striae; labrum bilobed; se. Brazil 

Llaqhastigmiis mmitanti Finnamore 

5 Mesosoma with at least pronotum red, sometimes entirely red 6 

5' Mesosoma black, at most pronotal lobe slightly red 7 

6 Transverse pronotal carina with prominent tooth at humeral angle (Figs. 70-72); median 
scutal groove and notauli extending over most of scutum (Fig. 71); Costa Rica, Colombia, 
Ecuador, Peru, Trinidad 19. pyrrhopyxis new species 

6' Transverse pronotal carina rounded at humeral angle, without tooth; no median scutal 
groove and notauli restricted to anterior third; Lesser Antilles: Dominica, Grenada, St. 
Vincent 22. thoracicus (Ashmead) 

7 Gena with tooth-like projection, ventrally (Fig. 12a); Peru, Ecuador, Colombia, Brazil 
(Mato Grosso) 6. hexagonalis (Fox) 

7' Gena smoothly rounded, without swellings, ventrally 8 

8 First metasomal tergum dull, densely microsculptured; se. Brazil 

18. pycuoglypticus new species 

8' First metasomal tergum shiny, without microsculpture 9 

9 Median scutal groove reduced, present only posteriorly and not reaching admedian lines 
(Figs. 8, 43) 10 

9' Median scutal groove well developed, reaching and often contiguous with admedian 

lines. (Figs. 3, 15, 79) 14 



16 Journal of Hymenoptera Research 

10 Pronotal lobes white 11 

10' Pronotal lobes black, occasionally brown or yellow-brown 12 

11 Pronotal lobe rounded (Figs. 42, 44); Central America 13. mystaxalbus new species 

11' Pronotal lobe toothed; South America 24. urqicus new species 

12 Transxerse pronotal carina with tooth at humeral angle larger than tooth on vertical 
pronotal carina; southern Mexico (Quintana Roo) to Panama .... 25. zephynis new species 

12' Transverse pronotal carina with tooth at humeral angle smaller than tooth on vertical 
carina; South America 13 

13 Vertex, posterior to micropore field, with shallow elliptical depression defined posteriorly 
by weak carina; flagellomeres II-Vl or more elongate, depressed basally, each with a 
broad shiny tylus on apical half; Ecuador, Peru, Bolivia, Brazil (Pernambuco) 

12. mauracis new species 

13' Vertex, posterior to micropore field, flat, without depression; flagellomeres cylindrical, 

not depressed basally; Venezuela, Ecuador 5. chincha new species 

14 Flagellomeres elongate, about 2x width, without tyli (Fig. 77); flagellomere XI broadly 
curved, more than 2x width; notauli extending -/, length of scutum (Fig. 79); se. Brazil 

21. snnicenis new species 

14' Flagellomeres relatively short, length subequal to width or, if elongate, then rarely 2x 
width and flagellomere XI cylindrical, not curved; tyli often present (Fig. 1); notauli often 
approaching posterior margin of scutimi 15 

15 Hyperstemaulus much narrower than scrobal sulcus; raised, linear tyli on flagellomeres 

III or IV to VII; Brazil (Ceara) 3. ccaraensis new species 

15' Hyperstemaulus equal to or wider than scrobal sulcus (measured vertically) (Fig. 16); 
flagellomeres usually without tyli, or tyli obscure 16 

16 Vertex, gena and posterior V, of scutum shiny, without microsculpture; median scutal 
groove tapered to a point just posterior to apex of admedian lines; Trinidad, Venezuela, 
Panama, Ct^lombia, Ecuador, Be^livia, Brazil (Goias) 17. prophorodontis new species 

16' Either vertex, gena or scutum dull, with microsculpture; median scutal groove reaching 

admedian lines 17 

17 Pronotal lobe toothed in dorsal view, forming a sharp acute angle, or flattened and wing- 
like in frontal view (Figs. 38-40); Colombia, to Bolivia, Brazil (Amazonas) 

11. kunkopteryx new species 

17' Pronotal lobe rounded in dorsal or frontal view, forming an obtuse angle (Figs. 15, 16) 

18 

18 Scutum striatopunctate on pcisterior -/,; Peru 2. caehikhus new species 

18' Scutum more or less smooth, without striae between grooves 19 

19 Flagellum with tyli (Fig. 29), or ventral brush of setae (Figs. 69, 81) 20 

19' Flagellum without tyli (at least not visible in profile) and without ventral brush of setae 

(Fig. 13); south of Guiana Shield to Brazilian Highlands 7. ictericoniis new species 

20 Flagellum with short, x^^ntral setal brush (Figs. 69, 81); tyli absent, flagellomere XI sym- 
metrical 21 

20' Flagellum without setal brush (Figs. 21, 29); tyli usually present, flagellomere XI often 

asymmetrical due to ventral tylus (Fig. 22) 23 

21 Mesosoma with at least pronotum red, sometimes entirely red; Costa Rica, Colombia, 
Ecuador, Peru, Trinidad 19. pyrrhopyxis new species 

21 ' Mesosoma black 22 

22 Vertex and frons uniformly microsculptured, dull; scutum with irregular striae on pos- 
terior -/v Peru 2. caeliikhus new species 

22' Vertex, anterior to mid ocellus, shiny (Fig. 82), microsculpture denser on frons than on 
vertex; scutum with striae usually less developed and only between notauli; se. Brazil, 
Paraguay, Peru Ecuador, Ccilombia, Venezuela, Suriname ... 23. trichodocems new species 

23 Flagellomere XI unmodified, cylindrical, without tylus (Fig. 30); Peru, Bolivia, Paraguay, 
Brazil (Bahia, Goias, Wlato Grosso, Sao Paulo) 10. iphis new species 



Volume 11, Number 1, 2002 17 

23' Flagellomere XI with tylus on ventral surface imparting an asymmetrical shape (Fig. 22); 

Colombia, Ecuador, Peru, Bolivia, Brazil (Mato Grosso, Para) 9. inti Finnamore 

24 Tyli of flagellomeres I-XI with sparse, short brush of setae (Figs. 69, 81); Argentina, Bo- 
livia, Brazil (Sao Paulo) 16. parauicus new species 

24' Tyli of flagellomeres without setal brush (Fig. 57), at mcost a few setae at apices of tyli 

(Fig. 49) 25 

25 Propodeum irregularly striate, without areolae; Venezuela 

20. strcpsiliiieatiis new species 

25' Propodeum areolate (Fig. 59) 26 

26 Pronotal lobe with carinate tooth or peg-like projection (Figs. 50-52) 27 

26' Pronotal lobe rounded, if somewhat pointed then not carinate (Figs. 59,60) 28 

27 Pronotal lobe toothed (acutely produced), white (Figs. 50-52); Venezuela, Colombia, Bo- 
livia, Brazil (Mato Grosso, Minas Gerais, Pernambuco, Rio de Janeiro) 

14. mytioy new species 

27' Pronotal lobe peg-like (bluntly produced), brown; Peru 4. ccromus new species 

28 Flagellomere XI with tylus imparting an asymmetrical shape; Argentina, Bolivia, se. Brazil 

1. aylaxiter new species 

28' flagellomere XI cylindrical, symmetrical, with at most indistinct tylus (Fig. 57); Mexico to 

Argentina 15. neotropictis (Kohl) 

29 Median clypeal lobe with 2 elongate setae arising from subapical semicircular depression 
(Figs. 18, 37, 38, 85, 86) ' " 30 

29' Median clvpeal lobe with 2 elongate setae arising from 2 narrowly separated subapical 
pits or a broad transverse depression (Figs. 10, 46, 62, 74) 33 

30 Mesopleuron shiny, hypoepimeral area unsculptured on ventral half or more, dorsal half 
finely microsculptured (Fig. 88); pronotal lobe rounded, conical (Fig. 87); scutum shiny 
posteriorly, often with short longitudinal irregular grooves (Figs. 87, 88); stigma of fore 
wing orange, brown or black; se. Brazil, Paraguay, Peru, Ecuador, Colombia, Venezuela, 
Suriname 23. trichoiiocerus new species 

30' Mesopleuron dull, microsculptured throughout (Fig. 20), OR pronotal lobe dorsally flat- 
tened (not conical), somewhat carinate anteriorly and laterally, OR posterior -/ -, of scutum 
with coarse regular striae; stigma of fore wing brown or black 31 

31 Scutum multistriate on posterior V,; mesopleuron shiny in part; pronotal lobe conical, 
not flattened or carinate; Peru 2. caclukhns new species 

31' Scutum without multiple striae except occasionally along posterior margin (Figs. 19, 39); 
mesopleuron microsculptured throughout OR pronotal lobe dorsally flattened, somewliat 
carinate anteriorly and laterally (Figs. 39-40) 32 

32 Pronotal lobe toothed, flattened dorsally, somewhat carinate anteriorly and laterally (Figs. 
38—40); hypoepimeral area shiny (Fig. 40); Colombia to Bolivia, Brazil (Amazonas) 

11. kimkopten/x new species 

32' Pronotal lobe rounded, conical, without carinae (Figs. 19-20); hypoepimeral area dull, 
microsculptured throughout (Fig. 20); south of Guiana shield to Brazilian Highlands .... 
7. ictericornis new species 

33 Median scutal groove absent, indistinguishable from other grooves along posterior mar- 
gin (Fig. 47) ! '. 34 

33' Median scutal groove present, at least as an elongate pit on posterior margin (Figs. 27, 

63) 36 

34 Clypeus white or yellow in apical third; mesosoma and petiole black; Central America 

13. mystaxalbus new species 

34' Clypeus black; mesosoma and petiole black to extensively red 35 

35 Median clypeal lobe absent; mesosoma and petiole usually extensively red; pronotal lobe 
white; Lesser Antilles: Dominica, Grenada, St. Vincent 22. thoraciciis (Ashmead) 

35' Median clypeal lobe present; mesosoma and petiole black, sometimes partially red on 



18 Journal of HYMENorxERA Research 

pronotum, scutum, scutellum, and mesopleuron; pronotal lobe yellow to red; southern 
Mexico (Quintana Roo) to Panama 25. zepln/nis new species 

36 Gena with ventral tooth (Fig. 12a); Brazil (Mato Grosso), Peru, Ecuador, Colombia 

6. hexagonalis (Fox) 

36' Gena smoothly rounded ventrally, at most with small swelling 37 

37 Scutum red 38 

37' Scutum all black, or red posteriorly 39 

38 Clypeus lateral to median lobe, simple, without tooth (Fig. 74); Costa Rica, Colombia, 
Ecuador, Peru, Trinidad 19, pyrrhopyxis new species 

38' Clypeus quadridentate, median lobe emarginate, bilobed, and flanked by small lateral 

tooth; Panama 8. ignithorax new species 

39 First metasomal tergvim dull, with dense, coarse microsculpture; se. Brazil 

18. pyciioglypticus new species 

39' First metasomal tergum shiny, without or with little microsculpture 40 

40 Pronotal lobe red, brown or black 41 

40' Pronotal lobe white 44 

41 Gena ventrally with small, subconical swelling near hvpostc^mal carina; Trinidad, Venezuela, 
Panama, Colombia, Ecuador, Bolivia, Brazil (Goias) 17. propJwrodoutis new species 

41' Gena ventrally flat, without swelling near hypostomal carina 42 

42 Lateral clypeal tooth located beneath antennal socket and separated from median lobe 

by deep emargination (Fig. 6); Venezuela, Ecuador 5. cliinclia new species 

42' Lateral clypeal tooth, if present, forming part of the median lobe, not distinct and not 
separated by emargination, appearing as a small basolateral angle on the median lobe 
43 

43 Pronotal lobe black or dark brown-black; lateral clypeal tooth present laterad of median 
clypeal lobe; Ecuador, Peru, Bolivia, Brazil (Pernambuco) 12. mauracis new species 

43' Pronotal lobe yellow-brown; lateral clypeal tooth absent; Brazil (Minas Gerais) 

24. iirqiciis new species 

44 Pronotal lobe toothed, peg-like or sharply acute (Figs. 38-40, 50-52) 45 

44' Pronotal lobe rounded (Figs. 25, 27, 28, 35, 36) 47 

45 Pronotal lobe with peg-like projection; median scutal groove reaching admedian lines; 
notauli often elongate, attenuating near posterior scutal margin; Peru 

4. ceromiis new species 

45' Pronotal lobe acute or dorsally flattened, wing-like; scutal grooves variable (Fig. 55) ... 4b 

46 Median scutal grtiove reaching admedian lines; notauli elongate (Fig. 39); pronotal lobes 
sometimes dorsally flattened (Fig. 38); Peru, Colombia 11. kiiiikopten/x new species 

46' Median scutal groove and notauli short, incomplete (Fig. 55); pronotal lobe conical, acute; 
Venezuela, Colombia, Bolivia, Brazil (Mato Grosso, Minas Gerais, Pernambuco, Rio de 
Janeiro) 14. niytior new species 

47 Scutum with 3 complete grooves, notauli reaching posterior scutal margin (Fig. 27); scu- 
tum shiny, microsculptured usually along anterior margin only 48 

47' Scutum with at most median groove complete (Fig. 67), notauli short; scutum usually 

microsculptured 49 

48 Vertex anterolaterally to mid ocellus shiny, without microsculpture (Fig. 25); Colombia, 
Ecuador, Peru, Bolivia, Brazil (Mato Grosso, Para) 9. inti Finnamore 

48' Vertex anterolaterally to mid ocellus dull, microsculptured (Fig. 33); Peru, Bolivia, Para- 
guay, Brazil (Bahia, Goias, Mato Grosso, Sao Paulo) 10. ipihis new species 

49 Lower hypoepimeral area shiny, without microsculpture; mid mesopleural area shiny and 
relatively large due to moderate and small size of foveae of scrobal sulcus, hyperster- 
naulus and omaulus (Fig. 68); occipital carina not raised ventrally (Fig. 68); antenna be- 
yond flagellomere II blackened, apical flagellomeres black; clypeal setae sparse, not ob- 
scuring underlying sculpture and clypeus punctures widely scattered (Fig. 66); Argentina, 
Bolivia, Brazil (Sao Paulo) 16, pnmnicus new species 



Volume 11, Number 1, 2002 



19 



49' Lower hypoepimeral area and mid mesopleural area microsculptured or shiny, the latter 
relatively small due to comparatively large size of foveae of scrobal sulcus, hyperster- 
naulus and omaulus (Fig. 64); occipital carina often slightly raised ventrally (Fig. 64); 
antenna usvially yellow to brown, seldom blackened; clypeal setae variable, scattered and 
not obscuring underlying microsculpture, to dense and obscuring underlying sculpture 
(Fig. 62); Mexico to Argentina 30 

50 Head and mesosoma extensixely microsculptured, usually entirely; scutum punctate, of- 
ten obscurely; clypeal setae sparse, not obscuring sculpture; Argentina, Bolivia, se. Brazil 

1. aylaxiter new species 

50' Vertex adjacent to ocelli, scutum, lower hypoepimeral area and mesopleuron often shin- 
ing (Fig. 64); scutum punctate to coarsely striatopunctate (Fig. 63); clypeal sculpture often 
obscured by dense appressed setae (Fig. 62) 51 

51 Micropore field of vertex small, elongate-triangular, usually much Ic^nger than wide, 
width about '/, ocellocular distance or less (Fig. 61); clypeus usually concealed by dense 
appressed setae (Fig. 62); USA (Texas) to Argentina 15. neotropiciis (Kohl) 

51' Micropore patch of vertex large, circular, extending about half ocellocular distance; clyp- 
eus with sparse setae that dc^ not obscure sculpture; se. Brazil . . 21. sunicerus new species 



1. lucastigmiis aylaxiter Finnamore 
new species 

Derivation of Name. — The species epithet 
is derived from two Greek words, aylax, 
meaning groove, and iter, meaning pas- 
sage, in reference to the meciian scutal 
groove found in this species. 

Diagnosis. — Males of aylaxiter can be rec- 
ognized on the basis of the narrow linear 
micropore field between the compound 
eye and lateral ocellus, flagellomeres with- 
out a ventral setal brush, tylus on flagel- 
lomere XI imparting an asymmetrical ap- 
pearance, and the rounded pronotal lobe. 
Females are difficult to recognize, but the 
following combination of characters 
should prove useful: median clypeal lobe 
with 2 narrowly separated subapical pits, 
clypeus not obscured by appressed setae, 
vertex microsculptured throughout, scu- 
tum with median groove not reaching ad- 
median lines, notauli short and not reach- 
ing scutal midlength, pronotal lobe refund- 
ed and white, hypoepimeral area micro- 
sculptured throughout, and metasomal 
tergum 1 shiny. This species most resem- 
bles paranicus and neotropvciis. Males are 
easily distinguished from other species us- 
ing the diagnostic characters. Females of 
aylaxiter are separated from parauieus on 



the basis of the entirely microsculpturecH 
hypoepimeral area (in parauieus the lower 
hypoepimeral area is shiny), and separat- 
ed from iieotropieus on the basis of the less 
extensively setose clypeus (clypeus usu- 
ally obscured by relatively dense ap- 
pressed setae in neotropieus) and by its dis- 
tribution which is apparently restricted to 
the Southern half of South America, 
whereas neotropieus ranges from Argenti- 
na to USA (Texas). 

Male. — Length 3.5-4.0 mm. Head. Fla- 
gellomeres without ventral brush of setae; 
narrow linear tyli present on flagellomeres 
I to XI, tylus on flagellomere XI imparting 
asymmetrical appearance and truncate tip; 
flagellomere I length 1.3 x maximum 
width as measured with tylus in profile; 
flagellomere X length 1.1 X maximum 
width as measured with tylus in profile. 
Clypeus obscured by dense appressed se- 
tae which extend up frons along inner 
margins of eyes to slightly less than height 
of scape; frons, vertex and gena entirely 
microsculptured; punctures of frons 
sparse, irregular, 2 or more diameters 
apart; punctures of gena obscured, more 
regular in ventral region, 3 or more di- 
ameters apart, without ventral tooth or 
swelling; narrow linear micropore field 
present between compound eye and lat- 



20 



Journal of Hymenoptera Research 



eral ocellus, without depression behind it; 
OOD 1.4 X LOD. Mesosoma. Transverse 
pronotal carina forming a right angle at 
humeral angle, toothed ventrally; trans- 
verse pronotal groove longitudinally stri- 
ate; pronotal lobe rounded, with weak an- 
terior carina; lateral pronotal area longi- 
tudinally striate. Scutum uniformly n^ii- 
crosculptured with sparse, irregular 
punctures, 2 or more diameters apart in 
mid region; median scutal groove extend- 
ing to scutal midlength, but not reaching 
admedian lines; notauli attenuated near 
scutal niidlength; posterior margin of scu- 
tum with a series of short irregular striae. 
Scutellum microsculptured with faint nie- 
dian groove and several punctures in lat- 
eral area. Setae of preomaular area sparse, 
not obscuring sculpture. Mesopleuron mi- 
crosculptured, punctures not evident; hy- 
persternaulus, scrobal sulcus, and on"iau- 
lus foveolate. Metapleuron microsculp- 
tured with longitudinal striae along pos- 
terior margin. Propodeum shiny, with 
weak microsculpture, areolate, except for 
shiny, partially striate region adjacent to 
metapleuron; propodeal enclosure defined 
by raised carina separating it from the lat- 
eral sphere. Metasoma. First tergum 
shiny, without microsculpture, with mi- 
nute, sparse punctures; succeeding terga 
with an oily sheen, with relatively larger 
punctures, 5 or more diameters apart. 
Sterna shiny, with oily sheen, with sparse 
punctures on basal sterna, but reaching 
greatest density on sternum VI where they 
are about 1 diameter apart. Color. Black. 
White: mandible, basal half; pronotal lobe. 
Yellow-brown: palpi; mandible, except 
base and apex; antenna, except flagellom- 
eres Vl-XI; tegula; fore leg, except coxa 
and femur; mid leg, except coxa and fe- 
mur; hind trochanter and hind tarsus. 

Female. — Length 4.0-5.0 mm. Similar to 
male except as follows: flagellomeres 
without tyli or specialized setae; flagel- 
lomere I length 1.7 X maximum width; 
clypeus shiny, setae more dense than in 
other species, but not obscuring sculpture. 



with punctures on median area 1 to 2 di- 
ameters apart; median clypeal lobe with 2 
teeth separated bv narrow emargination 
and a pair of narrowly separated subapi- 
cal pits; sculpture of frons along inner 
margin of eyes not obscured by appressed 
setae; micropore field present as an elon- 
gate triangle between compound eye and 
lateral ocellus; OOD 1.8 X LOD. 

Material Examined.— 7Q 6, 119 9. HO- 
LOTYPE MALE: Brazil: M.G. Ouro Preto 
lV-1954 N.L.H. Krauss (USNM). Para- 
types: ARGENTINA: Buenos Aires: 
Buenos Aires: 18-IV-1912 J.B. {16 MACN); 
4-V-1912 J.B. (19 MACN); 2-V-1915 J.B. 
(1(5 MACN); Moreno, Fritz (2 9 IIES). 
Punta Lara 26-1-1968 H. & M. Townes {IS 
AEIC). Catamarca: La Merced 26-VIII- 
1968 L. Pena (Id AEIC). Palo Labrado 27- 
111-1971 Fidalgo {16 IMLA). Cordoba/Ca- 
tamarca: Cordoba, Copacabana Fritz {26 
IIES). Jujuy: Ledesma Fritz {16 IIES). Per- 
ico del Carmen 21-X-1968 L. Pena {A 6 
CNCI). La Rioja: Cuesta de Miranda 
2020m 13-XI1-1971 Stange— Porter {16 
IMLA). Salta: Cachi 20-22-1-1966 C. Porter 
(1 6 MCZC). Magdalena 23-1-1966 H. & M. 
Townes (1 9 AEIC). Oran, Abra Grande 
18-25-X-1968 C. Porter {16 MCZC). Poci- 
tos Xn-1972 Fritz (Id IIES); XII-1971 Fritz 
(Id IIES). Rosario Lerma X-1984 Fritz (3d 
IIES). Tartagal A. Martines (Id IIES). Tuc- 
uman: Amaicha del Valle: 28-XII-1965 H. 
& M. Townes (Id AEIC); 29-XII-1965 H. 
& M. Townes (2d AEIC); 9-in-1966 Gar- 
cia — Porter (Id MCZC). Cuidad Tucuman 
24-11-1952 A. Ogloblin (Id IIES); [no date] 
Ogloblin (Id ilES); M906 Vezenvi (19 
HNHM); Tucuman, Horco Molle 10-XI- 
1967 C. Porter (Id MCZC). BOLIVIA: (2 9 
NHMW). Rio Beni, [Rurrenabanque] X 
W.M. Mann, Mulford Bio. Expl. 1921-22 
(Id USNM). El Beni: Cochabamba 10-4- 
1957 (2 9 FSAG). BRAZIL: [Brazilien] Fritz 
Muller, coll. G. Mayr (19 NHMW). Ama- 
zonas: R. Japura 13-IX-1904 Ducke (19 
MPEG). Bahia: [Bim Fim] 21-XI-1907 (19 
CMNH). [Liagoa Feia] 9-VI-1908 (19 
CMNH). Maranhao: [Varanahao Codo] 



Volume 11, Number 1, 2002 



21 



17-VI-1901 Ducke (1$ MPEG). Minas 
Gerais: Barbecena 23-X-1905 Ducke (19 
MPEG). Parana: Campina Grande nr. Cur- 
itiba 10-11-1966 H.& M. Townes (1 9 
AEIC). Rio de Janeiro: Parque Nacional 
Serra da Bocaina, S.J. Barreiros: 4-7-XI- 
1967 1600m Alvarenga & Seabra (1(5 19 
AEIC); XI-1968 1650m Alvarenga & Seabra 
(Ic? 69 AEIC); 13-17-1-1969 1600m Porter 
& Garcia {16 MCZC); IV-1969 800m P.M. 
Oliviera (2 9 AEIC); XI-1969 800m F.M. 
Oliviera (19 AEIC). Rio de Janeiro (19 
CMNH). Rio de Janeiro, Theresopolis: 9- 
X-1923 W.S. Bristowe B.M. 1967-510 {16 
BMNH); 12-III-1966 H. & M. Townes (2 9 
AEIC). Santa Catarina: Nova Teutonia: 
27°11'B 52°23'L F. Plaiimann (various 
dates) 1935-1967 {106 10 9 BMNH, 26 
16 9 MCZC, 1^ 3 9 OSUO); 27°11'B 
52°23'L IX-1967 300-500m F. Plaumann 
(19 MCZC); 27°11'B 52°23'L VI-1968 300- 
500m F. Plaumann {16 MCZC). Sao Pau- 
lo: S. Bocaina III-1973 (2 9 PMAE). Sao 
Paulo 1928 Bury J. Gyorgy (1 9 HNHM); 
(various dates) 1968-1969 V.N. Alin {16 
179 USNM); (various dates) 1969-1982 
(21 c5 419 ZMUM). 

2. Incastigmus caelukhus Finnamore 
new species 

Derivation of Name. — The name caelukhus 
is derived from two words, the Latin caelo, 
meaning to engrave, and the Quichuan 
term, ukini, meaning body, in reference to 
the striatopunctate scutum of this species. 

Diagnosis. — Males of this species can be 
recognized by the irregularly ridged, stria- 
topunctate scutum with complete or near- 
ly complete median scutal groove and no- 
tauli, the uniformly microsculptured frons 
and vertex, and the ventral brush of setae 
on the flagellomeres. Females can also be 
recognized by the ridged, striatopunctate 
sculpture of the scutum, the uniformly mi- 
crosculptured frons and vertex, and the 
semicircular depression on the median 
clypeal lobe from which long setae arise. 

Mrt/t'.— Length 3.5-4.0 mm. Head. Fla- 
gellomeres with a xentral brush of short 



fine setae, tyli absent; flagellomere I length 
1.8 X apical width; flagellomere X length 
1.2 X apical width; flagellomere XI 
straight, cylindrical, apex conical. Clypeus 
obscured by dense appressed setae which 
extend up frons along inner eye margin to 
height of antennal socket; frons and vertex 
dull, microsculptured, with sparse, irreg- 
ular, punctures, 3 or more diameters 
apart; gena microsculptured, obscurely 
punctate, without ventral tooth or swell- 
ing; micropore field present between com- 
pound eye and lateral ocellus, without de- 
pression behind it; lateral ocelli closer to 
each other than to eyes; OOD 1.3 x LOD. 
Mesosoma. Transverse pronotal carina 
entiing in a right angle at humeral angle, 
toothed ventrally; transverse groove lon- 
gitudinally striate; pronotal lobe rounded 
without anterior carina; lateral pronotal 
area longitudinally striate. Scutum ridged, 
striatopunctate, less so on lateral areas, 
microsculptured on anterior third, other- 
wise shiny; median scutal groove contig- 
uous with admedian lines, notauli contin- 
uous to posterior margin. Scutellum nii- 
crosculptured, with median sulcus, and 
scattered lateral punctures. Preomaular 
area anteriorly with sparse setae, sculp- 
ture visible. Mesopleuron with weak mi- 
crosculpture, mostly shiny; hypersternau- 
lus, scrobal sulcus, and omaulus, coarsely 
foveolate. Metapleuron shiny, weakly mi- 
crosculptured, with short longitudinal 
striae along posterior margin. Propodeum 
shiny, coarsely areolate, except area adja- 
cent to metapleuron which is shiny, un- 
sculptured; propodeal enclosure not dif- 
ferentiated from lateral spheres. Metaso- 
MA. Terga shiny with an oily sheen, with 
minute, obscure, punctures; sterna shiny, 
sparsely punctate, with punctures not in- 
creasing in density on posterior sterna. 
Color. Black. White: mandible, except 
apex; pronotal lobe. Yellow-brown: palpi; 
antenna; tegula; fore leg, except coxa; mid 
leg, except coxa; hind trochanter and tar- 
sus. 

Female. — Length 4.0-4.5 mm. Similar to 



22 



Journal of Hymenoptera Research 



male except as follows: flagellomere 1 
length 1.3 X apical width. Clypeus shiny, 
sparsely punctate and sparsely setose; me- 
dian clypeal lobe truncate, without teeth, 
with a large subapical semicircular de- 
pression froni which a pair of long setae 
arise; frons along inner eye margin ob- 
scured by appressed setae; OOD 1.7 X 
LOD; color as in male, except 1 specimen 
which has a red prothorax. 

Material Examined.— 3 d, 2 9. HOLO- 
TYPE FEMALE: Peru: 10km S. Chiclayo 
19-111-1951 Ross & Michelbacher (CASC). 
Paratypes: PERU: 10km S. [Chiclayo] 19- 
111-1951 Ross & Michelbacher (2(5 CASC). 
Lambayeque: 18km E. Olmos 22-V11-1982 
R.B. Miller, L.A. Stange (19 FSCA). Piura: 
Piura l-XI-1910-7-1-1911 C.H. Townsend, 
Leber {\6 USNM). 

3. Incastigmus cearaensis Finnamore 
new species 

Figs. 1-4 

Derivation of Name. — Cearaensis is named 
in reference to the state of Ceara in Brazil, 
from where most of the specimens in this 
study have been collected. 

Diagnosis. — The narrow hypersternau- 
lus and the presence of tyli on flagellom- 
eres 111 to V will distinguish the male of 
this species from others in the genus. The 
female is unknown. 

M/?/t'.— Length 3.5-4.0 mm. Head. Fla- 
gellomeres without specialized setae; lin- 
ear tyli present on flagellomeres 1 to V or 
VII; flagellomere I length 1.5 X apical 
width; flagellomere X length 1.3 X apical 
width; flagellomere XI straight, cylindri- 
cal, apex conical. Clypeus obscured by 
dense appressed setae which extend up 
frons along inner margin of eye to about 
height of antennal socket; head entirely 
microsculptured, slightly less so on vertex; 
punctures strong, sparse, three or more di- 
ameters apart on vertex; gena microsculp- 
tured, with punctures in lower area 3 or 
more diameters apart, without ventral 
tooth or swelling; small oval micropore 
field evident between compound eye and 



lateral ocellus; without depression behind 
it; lateral ocelli slightly closer to eyes than 
to each other; OOD 1.4 x LOD. Mesoso- 
MA. Transverse pronotal carina toothed at 
humeral angle, and toothed ventrally; 
transverse pronotal groove longitudinally 
striate; pronotal lobe rounded, without an- 
terior carina; lateral pronotal area longi- 
tudinally striate. Scutum microsculptured, 
dull, with several longitudinal striae along 
posterior margin; notauli attenuated near 
scutal midlength; median scutal groove 
present posteriorly, attenuated near scutal 
midlength; scutal punctures strong, 
sparse, 1 or more diameters apart. Scutel- 
lum microsculptured, with several punc- 
tures in median lateral area. Preomaular 
area anteriorly with sculpture visible and 
sparse setae. Mesopleuron microsculp- 
tured, dull, with sparse, obscure, punc- 
tures; hypersternaulus narrow, linear, 
with relatively fine foveae; scrobal sulcus, 
and omaulus coarsely foveolate; meta- 
pleuron dull, microsculptured, with sev- 
eral short longitudinal striae along poste- 
rior margin. Propodeum shiny, with weak 
microsculpture, coarsely areolate, except 
area adjacent to metapleuron which has 
fine microstriate sculpture; propodeal en- 
closure not differentiated from lateral 
spheres. Metasoma. Tergum I shiny with 
weak microsculpture; succeeding terga 
with oily sheen, punctures sparse, ob- 
scure, many diameters apart. Sterna shiny 
with weak microsculpture imparting an 
oily sheen, punctures obscure, sparse, 3 or 
more diameters apart. Color. Black. 
White: pronotal lobe. Yellow-brown: pal- 
pi; mandible, except apex; antenna; fore 
leg, except coxa; mid leg, except coxa; 
hind tarsus; apical sterna. 

Female. — Unknown. 

Material Examined.— A 6. HOLOTYPE 
MALE: Brazil: SP, Faz. Campininas, Mogi 
Guacu 29-31 -XII-1969 JM & BA Campbell 
(CNCI). Paratypes: BRAZIL: Ceara: Ser- 
rado Araripe, Crato V-1969 M. Alvarenga 
(3(5 PMAE). 



Volume 11, Number 1, 2002 



23 




Figs. 1—4. Incastigiinis ccnraoisis 6.1, Mid flagellomeres of antenna. 2, Head, dorsal; arrow indicates micro- 
pore field. 3, Mesosoma, dorsal. 4, Meseisoma, lateral (tooth on mesopleuron is an artifact). 



4. Iticastigmiis ceromns Finnamore 
new species 

Deriz^ntion of Nnnic. — The species epithet 
is a derivation of two Greek words, kcros, 
meaning horn, and onios, meaning shoul- 
der, in reference to the peg-like pronotal 
lobe of this species. 

Diagnosis. — This species has the greatest 
development of the pronotal lobe of any 
species in the genus. The blunt, peg-like 
pronotal lobe in both males and females 
separates ccroiiiiis from all other liicastig- 



imis. In addition, the male pronotal lobe is 
brown, and the vertex has a linear micro- 
pore field between the compound eye and 
lateral ocellus; in the female the micropore 
field is an elongate triangle, the pronotal 
lobe is white, and notauli do not reach the 
posterior scutal margin, but attenuate on 
the posterior half of the scutum. 

Male. — Length 4.5 mm. Head. Flagel- 
lomeres setose throughout, but without 
specialized ventral brush of setae, at most 
with a few specialized setae on the apices 



24 



JOURNAI, OF HYMENOPTERA RESEARCH 



of the flagellomeres; all flagellomeres with 
linear tyli; tylus on flagellomere XI im- 
parting a slight asymmetrical appearance 
and blunt tip; flagellomere I length 1.2 X 
maximum width as measured with tylus 
in profile; flagellomere X length 1.2 X 
maximum width as measured with tylus 
in profile. Clypeus obscured by dense ap- 
pressed setae which extend up frons along 
inner margins of eyes to approximately 
half height of scape; frons coai"sely mi- 
crosculptured, somewhat shiny anterior to 
ocelli and on vertex; punctures of upper 
frons and vertex sparse, irregular, 3 or 
more diameters apart; vertex, posterior to 
ocelli, microstriate; gena microsculptured, 
longitudinally striate on lower half, with- 
out ventral tooth or swelling; linear micro- 
pore field present between compound eye 
and lateral ocellus; without depression be- 
hind it; OOD 1.7 X LOD. Mesosoma. 
Transverse pronotal carina toothed at hu- 
meral angle and toothed ventrally; trans- 
verse pronotal groove longitudinally stri- 
ate; pronotal lobe produced as blunt peg- 
like projection with strong anterior carina; 
lateral area of pronotum longitudinally 
striate. Scutum microsculptured, punc- 
tures small, sparse, irregular, 1 to many 
diameters apart in midregion; notauli ex- 
tending to near scutal midlength, attenu- 
ating on posterior half, not reaching pos- 
terior margin; median scutal groove atten- 
uating at admedian lines; posterior mar- 
gin of scutum with a series of ridges 
extending to posterior quarter of scutum. 
Scutellum microsculptured, with median 
longitudinal sulcus and sparse scattered 
punctures. Sculpture of preomaular area 
not obscured by setae. Mesopleuron with 
microsculpture in dorsal region, shiny, 
weakly microsculptured ventrally; hyper- 
sternaulus, scrobal sulcus, and omaulus 
coarsely foveolate. Metapleuron micro- 
sculptured, with several longitudinal stri- 
ae along posterior margin. Propodeum 
shiny, without niicrosculpture, areolate 
except small microstriate region adjacent 
to metapleuron; propodeal enclosure not 



differentiated from lateral spheres. Meta- 
SOMA. First tergum shiny with minute ob- 
scure punctures; succeeding terga with an 
oily sheen, punctures sparse, minute, ob- 
scure. Sterna shiny, with oily sheen, punc- 
tures sparse on basal sterna, not increasing 
in density until sternum VI where they are 
about 2 diameters apart in median region. 
Color. Black. Yellow-brown: palpi; man- 
dibles, except apex; antenna, except apical 
flagellomeres; tegula; pronotal lobe; fore 
leg, except coxa; mid leg, except coxa; 
hind tarsus; apical sternum and tergum. 

FcDinle. — Length 4.0-5.0 mm. Similar to 
male except as follows: flagellomeres 
without tyli or specialized setae; flagel- 
lomere I length 1.5 X maximum width; 
clypeus shiny, with weak microsculpture, 
punctures sparse in median region and 1 
or more diameters apart; median clypeal 
lobe with 2 blunt teeth separated by a 
shallow notch, and with 2 narrowly sep- 
arated subapical pits; sculpture of frons 
along inner margins of eyes not obscured 
by setae; upper frons and vertex more 
shiny than in male, microstriae not as ev- 
ident; gena more shiny than in male, mi- 
crosculpture weak ventrally, longitudinal 
striae on lower half present to absent; mi- 
cropore field present as an elongate tri- 
angle between compound eye and lateral 
ocellus; OOD 2.2 X LOD; scutum with 
weak microsculpture on posterior half, 
more shiny than in male; pronotal lobe 
white. 

Material Examined.— 1 6,7 9. HOLO- 
TYPE MALE: Peru: Carpish Mts. 40mi S. 
Tingo Maria 28-XII-1954 E.l. Schlinger & 
E.S. Ross (CASC). Paratypes: PERU:' Cuz- 
co: Rio Urubamba 3km above Machu Pic- 
chu 2050m 18-IV-1983 C. & M. Vardy B.M. 
1983-217 (19 BMNH). Machu Picchu mu- 
seum 1,385m 11-14-VI1I-1971 C. & M. Var- 
dy B.M. 1971-533 (1 9 BMNH). Agua Cal- 
liente 21-28-XII-1983 (19 PMAE). Huanu- 
co: Carpish Mts. 40mi S. Tingo Maria 28- 
XII-1954 E.I. Schlinger & E.S. Ross (39 
CASC). 26mi E. Tingo Maria lO-XII-1954 
1100m E.I. Schlinger & E.S. Ross (IJ 



Volume 11, Number 1, 2002 



25 




Figs. 5-8. //;a;s//v^/;;//> clunclui ?. 5, Head, dorsal. 6, Clypeus, oblique dorsal view. 7, Mesosoma, lateral. 8, 
Mesosoma, dorsal. 



CASC). Pasco: Oxapampa 2,200m 7-III- 
1979 M. Cooper B.M. 1979-216 (19 
BMNH). 

5. Incastigmtis chiuclm Finnamore 
new species 

Figs. 5-8 

Dcrivntioii of Name. — Chiiichn is a Que- 
chuan term, meaning northern in refer- 
ence to the distribution of this species in 
the northern part of South Anierica. 

Din;j^}iosis. — Males of this species can be 



recognized on the basis of the short me- 
dian scutal groove, the dark, toothed 
pronotal lobe, and the absence of a de- 
pression posterior to the micrc^pore field. 
Females can be recognized on the basis of 
the short median scutal groove, the dark, 
toothed pronotal lobe, and the acute lat- 
eral tooth on the apical margin of the clyp- 
eus. 

Mnlc. — Length 4.0 mm. Head. Flagel- 
lomeres without a brush of specialized 
setae or tyli; tlagellomere 1 length 1.6 X 



26 



Journal of Hymenoptera Research 



apical width; flagellomere X length 1.3 X 
apical width; flagellomere XI straight, 
cylindrical, apex conical. Clypeus ob- 
scured by dense appressed setae which 
extend up frons along inner margin of 
eyes to a point about twice the height of 
antennal socket. Frons microsculptured, 
punctures not evident; vertex shiny, 
with microsculpture and punctures scat- 
tered, sparse, many diameters part; gena 
microsculptured, more shiny than ver- 
tex, sparse punctures more evident ven- 
trally, without ventral tooth or swelling; 
lateral ocelli closer to each other than to 
compound eyes; OOD 1.5 x LOD. Me- 
SOSOMA. Transverse pronotal carina 
toothed at humeral angle and toothed 
ventrally, transverse pronotal groove 
longitudinally striate, pronotal lobe with 
anterior carina ending in acute tooth; lat- 
eral region of pronotun^i longitudinally 
striate. Scutuni microsculptured, some- 
what shiny; punctures coarse, sparse, 3 
or more diameters apart; notauli present 
anteriorly, attenuated posteriorly near 
scutal midlength, median groove present 
posteriorly, attenuated anteriorly, not 
reaching admedian lines; posterior mar- 
gin of scutum with a series of short ridg- 
es parallel to, but shorter than, median 
groove. Scutellum microsculptureci, with 
several punctures in median area, and a 
median longitudinal sulcus. Mesopleu- 
ron entirely microsculptured, apparently 
impunctate; preomaular area anteriorly 
with sculpture evident, setae sparse; hy- 
persternaulus, scrobal sulcus, and omau- 
lus coarsely foveolate. Metapleuron 
coarsely microsculptured with several 
longitudinal striae. Propodeum shiny, 
with weak microsculpture, coarsely are- 
olate throughout, except area adjacent to 
metapleuron which is longitudinally 
striate; propodeal enclosure not differ- 
entiated from lateral spheres. Metaso- 
MA. Tergum 1 shiny, without microsculp- 
ture, punctures sparse, obscure. Terga 
beyond first tergum with oily sheen. 
Sterna shiny, with weak microsculpture; 



punctures sparse, becoming increasingly 
dense on more posterior sterna; punc- 
tures of sternum VI about 1.5 diameters 
apart. Color. Black. Yellow-brown: pal- 
pi; mandible, except apically; antennae 
except apical flagellomeres; tegula; pron- 
otal lobe; fore leg, except coxa; mid leg, 
except coxa; hind tarsus, hind trochan- 
ter, and apex of hind tibia; sterna VI to 
VIII. 

Female. — Length 4.5-5.5 mm. Similar to 
male except as follows: flagellomere I 
length 1.8 X apical width; clypeus shiny 
with punctures sparse, more than 5 di- 
ameters apart medially; median clypeal 
lobe reduced to a narrowly rounded pro- 
trusion, median teeth not evident and me- 
dian lobe with a pair of subapical circular 
pits from which several long setae arise; 
acute lateral tooth present on clypeal mar- 
gin, situated below antennal socket; frons 
along inner eye margin not obscured by 
appressed setae. OOD 2.2 x LOD. Scutum 
and mesopleuron generally with less mi- 
crosculpture, more shiny. 

Materia] Examined.— 2 c? 13 9. HOLO- 
TYPE FEMALE: Venezuela: Aragua, 
Rancho Grande, Portachuelo 1100m 22- 
V-1981 J.L. Garcia & J. Clavijos (IZAV). 
Paratypes: ECUADOR: Pichincha: Tin- 
alandia 800m 11-1983 M. Sharkey & L. 
Masner (2 9 PMAE). 16km SE. Sto. Do- 
mingo 500m Tinalandia 4-14-VI-1976 S. 
& J. Peck (1 9 CNCI). VENEZUELA: Ar- 
agua: Rancho Grande, Portachuelo: 
1100m 21-V-1981 ].L. Garcia & J. Clavijos 
{16 29 IZAV); 1100m 22-V-1981 J.L. 
Garcia & J. Clavijos (3 9 IZAV). Bolivar: 
Kavanayen 20-X-1972 J. & B. Bechyne 
(19 IZAV). Falcon: [Cerrogalicia] 1500m 
3-XII-1971 J. & B. Bechyne (19 IZAV). 
Cerro Galicia 1500m 19-XI-1971 J. & B. 
Bechyne (19 IZAV). Lara: Cubiro 6-V- 
1981 H.K. Townes (1(5 AEIC). Merida: 
Valle Grande 23-VII-1988 C. Porter & L. 
Stange (1 9 FSCA). Trujillo: La Mesa 26- 
Vn-lt)66 I. & B. Bechyne (1 9 IZAV). 



Volume 11, Number 1, 2002 



27 



6. Incastigmtis hexagoualis (Fox) 
new combination 

Figs. 9-12 

Stigiiiiis hcxngoiinlis Fox 1897:379. Lectotype, fe- 
male CMNH. Brazil: Chapada, Dec; exam- 
ined. 

Dinguosis. — Both sexes can be easily rec- 
ognized by the tooth-like projection on the 
ventral gena near the hypostomal carina. 
Females can be distinguisheci from female 
pjrophowdoiitis, which also have a small 
tooth-like swelling on the lower gena, by 
the presence of a pair of teeth on the me- 
dian clypeal lobe; female pwplioiviloiitis 
have a truncate median clypeal lobe, with- 
out teeth. 

M^7/t'.— Length 3.5-4.2 mm. Head. Fla- 
gellon"ieres without tyli, but with a ventral 
brush of setae; flagellomere 1 length 2.0 X 
apical width; flagellomere X length 1.2 X 
apical width; flagellomere XI straight, cy- 
lindrical, apex conical. Frons and clypeus 
with uniform microsculpture; vertex and 
gena shiny, with sparse punctures sepa- 
rated by 5 or more puncture diameters. 
Clypeus obscured by dense setae which 
extend up frons along eye margin to 
height of antennal socket. Gena shiny, 
without microsculpture, sparsely punc- 
tate, not striate, and with prominent ven- 
tral tooth-like swelling. Micropore field 
present as a discrete oval patch on vertex 
between lateral ocellus and compound 
eye, without a depression behind it. Ocelli 
closer to each other than to eyes. OOD 1.6 
X LOD. Mesosoma. Transverse pronotal 
carina with prominent tooth at humeral 
angle and prominent ventral tooth; trans- 
verse pronotal groove with several longi- 
tudinal striae. Pronotal lobe tooth-like, 
conical. Pronotum laterally striate. Scutum 
shiny, with microsculpture anteriorly. No- 
tauli well developed, present on anterior 
half of scutum. Median groove of scutum 
well developed and complete, or some- 
times attenuated anteriorly between ad- 
median lines, broadened posteriorly. Pos- 
terior margin of scutum with several short 



parallel striae on each side of the median 
groove. Scutellum shiny, with weak mi- 
crosculpture, generally impunctate, but a 
small group of punctures present laterally. 
Mesopleuron shiny, without microsculp- 
ture, impunctate. Preomaular area anteri- 
orly with sparse setae. Hypersternaulus 
coarsely foveolate, often with only two 
large foveae. Scrobal sulcus and omaulus 
coarsely foveolate. Metapleuron weakly 
microsculptured on ventral half, other- 
wise shiny, impunctate. Propodeum 
shiny, without microsculpture; propodeal 
enclosure and lateral spheres not differ- 
entiated, coarsely areolate throughout, ex- 
cept basal area adjacent to metapleuron 
which is smooth and shiny, without sculp- 
ture. Metasoma. Terga shiny, without mi- 
crosculpture; punctures sparse, obscure. 
Sterna shiny; punctures sparse, weak. 
Color. Black. Yellow-brown: antenna, ex- 
cept dorsal apex of flagellum; mandible, 
except apex; palpi; tegula; fore and mid 
legs, except coxae; hind coxal apex, hind 
trochanter, hind tibia \^entrally, and hind 
tarsus. 

Feninlc. — Length 4.0-5.25 mm. Similar to 
male except as follows: flagellomere I 
length 2.25 X apical width. Clypeus shiny, 
without appressed setae, sparsely to 
densely punctate; median clypeal lobe 
with a pair of teeth separated by a deep 
U-shaped emargination; the base of each 
tooth bears a large pit from which arises 
an elongate seta. Lower gena with a prom- 
inent tooth-like swelling. OOD 2.4 X LOD. 
Occipital carina simple, not foveolate. Col- 
or as in male for the dark form, except 
pronotum, scutum, scutellum, and upper 
mesopleural area, orange-red in light 
form. 

Material Examined. — 3 6, 28 9. CO- 
LOMBIA: Caqueta: Yuruyaco, 73km SW. 
Florencia (BMNH). Meta: Rio Duda 
(BMNH). Putumayo: Moco. (BMNH); Mo- 
coa, 8mi S. Puerto Assis 350m (BMNH). 
Villa Garzon (BMNH). ECUADOR: Mo- 
rona: Macas, 6km E. Santiago, Cord. Cu- 
tucu (BMNH). Napo: Limoncocha (on Rio 



28 



Journal of Hymf-noptera Research 




Figs. 9-12. Iiicasti;^iiiii^ licxip^oiialis + . 9, Head, d(.)rsal. 10, Head, fnmtal. 1 I, Mesusmna, lateral. 12, Mesosoina, 
dorsal. 12a, Head, lateral, with arrow indicating genal tooth. 



Volume 11, Number 1, 2002 



29 



Napo) (19 FSCA). Misahualli (1 9 PMAE). 
Tena (1 9 PMAE). PERU: [Chanchamayo] 
(BMNH). Colonia: Junin (BMNH). La 
Merced, 18 mi NE. Perene, Rio Perene (1 9 
CASC). Huanuco: Tingo Maria (BMNH). 
Junin: Paratuchali (19 PMAE). Satipo 
(BMNH). Loreto: Iquitos, Barillal (19 
PMAE). BRAZIL: [Chapada] (3 9 CMNH). 
[Corumba] (19 CMNH). [Santarem] (19 
USNM). Mato Grosso: 12°50'S 5r47'W 
(BMNH). Cerrado (BMNH). 

7. Incastigmus ictericoniis Finnamore 
new species 

Figs. 13-20 

Dcrivntioii of Nauic. — The name ictcricor- 
nis is derived from the Greek ikterikos, 
meaning yellowish, and the Latin coniii, 
meaning horn, in reference to the yellow- 
ish antemia of this species. 

Diagnosis. — Females are easily distin- 
guished from other species by the follow- 
ing combination of characters: the median 
lobe of the clypeus has a subapical semi- 
circular depression with several long se- 
tae, the pronotal lobe is rounded, and the 
scuta 1 punctures are normal, not stria to- 
punctate. Males are not easily separated 
from other species, but the following char- 
acters will be of some assistance: the fla- 
gellum without ventral brush of setae or 
tyli, and flagellomeres I and X length less 
than 2 X apical width, the gena without 
ventral tooth or swelling, mesosoma 
black, pronotal lobe rounded, median 
groove of scutum reaching admedian 
lines, and tergum I of metasoma shiny, 
without microsculpture. 

Male. — Length 4.0-4.5 mm. Head. Fla- 
gellomeres without tyli or specialized se- 
tae ventrally; flagellomere I length 1.6 X 
apical width; flagellomere X length 1.3 X 
apical width; flagellomere XI straight, cy- 
lindrical, apex conical; clypeus obscured 
by dense appressed setae which extend up 
frons along inner eye margin to half the 
height of scape; frons inicrosculptured, 
punctures obscure; vertex shiny, weakly 
microsculptured, punctures irregular, 2 or 



more diameters apart; gena without ven- 
tral tooth or swelling, microsculptured, 
punctures obscure and scattered; micro- 
pore field present as an oval patch be- 
tween compound eye and ocellus, without 
depression behind it; lateral ocelli closer to 
each other than to compound eye; OOD 
1.4 X LOD. Mesosoma. Transverse pron- 
otal carina toothed at humeral angle, and 
toothed ventrally; transverse pronotal 
groove longitudinally striate; pronotal 
lobe rounded, without anterior carina; lat- 
eral region of pronotum longitudinally 
striate. Scutum microsculptured through- 
out, punctures irregular, 1 or more diam- 
eters apart; notauli usually attenuating 
about scutal midlength, in some speci- 
mens notauli reach posterior quarter of 
scutum; median scutal groove reaching 
admedian lines. Scutelluni microsculp- 
tured, with median longitudinal sulcus, 
and several punctures laterally. Preomau- 
lar area anteriorly with sparse setae that 
do not obscure sculpture. Mesopleuron 
microsculptured on hypoepimeral area, 
shiny with less microsculpture on ventral 
regions; hypersternaulus, scrobal sulcus, 
and omaulus coarsely foveolate; meso- 
pleuron microsculptured, with several 
short longitudinal striae along posterior 
margin. Propodeum shiny, weakly mi- 
crosculptured, coarsely areolate over most 
of its surface, except area adjacent to me- 
tapleuron which is shiny, without sculp- 
ture; propodeal enclosure not differenti- 
ated from lateral spheres. Metasoma. First 
tergum shiny, without microsculpture, 
succeeding terga with an oily sheen, 
sparsely punctate with punctures many 
diameters apart. Sterna similar in sculp- 
ture to terga with punctures slightly more 
dense on posterior sterna and 3 or more 
diameters apart in lateral regions. Color. 
Black. White: mandible, except apex; 
pronotal lobe. Yellow-brown: palpi; anten- 
nae; tegula; fore leg, except coxa and fe- 
mur; mid leg, except coxa and femur; hind 
tarsus. 

Fciiinlc. — Length 4.0-5.0 nim. Similar to 



30 



Journal of Hymenoptera Research 




Figs. 13-16. liicn^tlgniii^ ictcrlconiisu . 13, Mid flagellomeres of antenna. 14, Head, dorsal. 15, Mesosoma, 
dorsal. 16, Mesosoma, lateral. 



male except as follows: flagellomere 1 
length 1.8 X apical width; clypeus shiny, 
without microsculpture, or with weak 
microsculpture medially; clypeal punc- 
tures 1-3 diameters apart medially; me- 
dian clypeal lobe with 2 teeth separated 
by a shallow emargination, long setae 
arising from subapical semicircular de- 
pression. Inner eye margin sparsely se- 
tose with sculpture not obscured; OOD 
2.0 X LOD. 

Material ExaiiiiiWii.—2S 6,7 9. HOLO- 



TYPE MALE: Peru: Madre de Dios: Pto. 
Maldonado 1-11-1-1984 L. Huggert 
(PMAE). Paratypes: ARGENTINA: Salta: 
Oran, Abra Grande 18-25-X-1968 C. Por- 
ter (3c? MCZC); 18-1V-5-V-1969 C. Porter 
(1(5 MCZC). Tucuman: Horco Molle, nr. 
Tucuman 15-21-V-1966 L. Stange (IJ 
IMLA). BOLIVIA: La Paz: Cavinas, Rio 
Beni VII W.M. Mann Mulford Bio. Expl. 
1921-22 (19 USNM). BRAZIL: Amazon- 
as: [Jeffe] 7-IX-1904 Ducke {\6 MPEG). 
Bahia: Itabuna CEPEC XI-1978 P.P. Ben- 



Volume 11, Number 1, 2002 



31 



ton {IS BMNH). Espirito Santo: Unbar- 
es, XI-1967 F.M. Oliveira {16 PMAE). Mi- 
nas Gerais: Aguas Vermelhas 15°45'S 
41°28'W 800m XII-1983 Alvarenga (Id 
AEIC). Barbacena 26-X-1905 Ducke (Id 
MPEG). S. Caraca, S Barbara III-1971 F.M. 
Oliveira (2$ PMAE). Rio de Janeiro: Re- 
presa Rio Grande, Guanabara V-1972 M. 
Alvarenga {26 19 PMA). Rio de Janeiro 
VIII {26 USNM). Teresopolis 9-III-1966 
H. & M. Townes (19 AEIC). Rondonia: 
[Ft.] Principe [da Beira?] Rio Guapore {16 
CMNH). Santa Catarina: Nova Teutonia, 
1-1968 F. Plaumann (1 9 MCZC). Sao Pau- 
lo: Sao Paulo {\6 ZMUM). ECUADOR: 
[Bucay] 1000' 7-X-1922 F.X. Williams {16 
BPBM). PARAGUAY: [no locality] Fie- 
brig {16 HNHM). Cordillera: Col. [Co- 
lonia] Piareta XII-1971 Pena {26 IIES). 
PERU: Huanuco: Tingo Maria 1km E. 2- 
VIII-1971 2000' P.S.&H.L. Broomfield 
B.M. 1971-486 (19 BMNH). Madre de 
Dios: Pto. M